November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorininae 417 J - CY e—- C 7 Ly / As) THE FAMILY LITTORINIDAE IN THE INDO-PACIFIC Part I. The Subfamily Littorininae by JOSEPH ROSEWATER Division of Mollusks National Museum of Natural History Washington, D.C. 20560 U.S.A. Abstract The classification of tropical Indo-Pacific Lit- torinidae (Mollusca: Mesogastropoda: Lit- torininae) has been revised. Thirty four Recent and six Tertiary fossil species belonging to six generic groups are described, and/or redescribed, and figured. Complete systematic synonymies are given, together with discussions of relationships, biology and zoogeography. One new subgenus and two new subspecies are described; one new name is assigned. Spawn characteristics of Lit- torinidae are reviewed. A list is given of recog- nized taxa of world-wide Littorinidae belonging to the subfamily Littorininae. Indo-Pacific Littorinidae Representatives of the family Littorinidae are found in most regions of the world occupying habitats from relatively shallow waters below the intertidal zone to situations high above the sea where they may be wet only occasionally by spray. Certain species are unusual among marine gastropods in their ability to survive for long periods without immersion in sea water (Rose- water, 1963a). These species may be considered as living examples of some stages in the invasion of the land by marine snails. The genus Crem- noconchus Blanford, 1869, inhabits fresh water, although it is doubtful that it invaded that habitat from the sea. Although the present paper is main- ly concerned with marine Indo-Pacific Lit- torinidae, the list of recognized taxa on page 423 [05-267] contains the names of all valid species recognized by me in the various generic groups. Family Characters Littorinidae are generalized mesogastropods, and members of the type-genus, Littorina, have nothing outstanding about their appearance. It is, perhaps, this generalized condition which helps to distinguish them from other closely related gastropod families which possess at least some prominent characteristics. “Typical” Littorina, best exemplified by the type-species Littorina littorea (Linné, 1758) (see pl. 326, figs. 1, 2) have thick-walled turbinate shells, are usually non- umbilicate, and have paucispiral opercula, al- though members of the genus Echininus disre- gard the general rule and have umbilicate shells and multispiral opercula. Sculpture varies from nearly smooth to spirally striate, axially furrowed, or nodulose. Some species of Tectarius are quite elaborately sculptured. The anatomy of Littorina was described in some detail by Fretter and Graham (1962, see our pl. 327, figs. A and B). There is a moderately- sized, ditaxic foot bearing the operculum on its dorsal posterior surface; a head with two tentacles having dark eyes at their outer bases; a central, anterior snout with the mouth at its extremity. All species in the family so far as is known are dioecious; the male bears a penis and the female has a well-marked groove on the side of the “neck” for passage of eggs. Fertilization is in- ternal, and pelagic development is the rule, al- though several species have evolved ovovivipari- ty. Species differences may be noted in penile anatomy, reproductive habits and characters of egg capsules (see table). [05-261] 418 Littorinidae Joseph Rosewater Littorinidae ayaa gai Cee y ii}; ; 1 i oii Plate 325. Subfamily Littorininae (explanation on opposite page). (all figures about natural size) [05-262] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorininae 419 Littorinidae most often are confused with such families as Planaxidae. The resemblance between the two is considerable but superficial. This con- fusion may be dispelled if it is remembered that the columella of Planaxidae terminates in a short, “pinched” siphon, while that of Littorinidae is always simple. The closest marine relative, Lacu- nidae, is readily recognized, due to the omnipre- sent umbilical chink, usually more conical shells, and metapodial tentacles. Details of habitat and reproductive characters are also different. Other possible Littorina “look-alikes” may be found among the Fossaridae and Trochacea and also in certain land and freshwater prosobranch families, such as the Helicinidae and Hydrobiidae. The latter do not, of course, occur in a habitat similar to that occupied by Littorinidae. Trochacea differ significantly in having the interior of the shell nacreous and also in being quite different ana- tomically. Throughout the descriptions of the species, measurements of shell size are given in both millimeters and inches. The maximum length to which shells grow is given. The factor of obesity was determined by dividing the width of the shell by the length. Spawn Characteristics A review of the literature on reproduction in Littorinidae reveals that most species, about which the information is known, release pelagic capsules. A few deposit egg masses on a suitable substrate and produce living young. Below is a partial list of species, arranged according to spawn type and faunal region. Following each species I have given the references and, where necessary, the correct name in square brackets. A. Pelagic Capsule— Indo-Pacific: Littorina brevicula: Kojima (1957, 1958a); Habe (1958); Yamamoto and Habe (1962); Hirai (1963); Yamaguchi (1967). L. coccinea (per- sonal observations, see under Description of L. coccinea). L. pintado: Ostergaard (1950); Whip- ple (1965); Struhsaker (1966). L. strigata [= L. undulata?]: Kojima (1958c); L. undulata (per- sonal observations, see Description of L. undulata). Nodilittorina pyramidalis: Kojima (1958b). Gran- ulilittorina granularis [= G. millegrana]: Tokioka and Habe (1953); Habe (1955); Yamamoto and Habe (1962); Hirai (1963). G. picta: Whipple (1965); Struhsaker (1966, 1968b). North Pacific: Littorina squalida: Habe (1958); Kojima (1958b, e); Yamamoto and Habe (1962). Eastern Pacific: Littorina planaxis: MacGinitie and MacGinitie (1949), Gibson (1964). Western and Eastern Atlantic: Littorina lit- torea: Thorson (1946). Eastern Atlantic: Littorina neritoides: Lebour (1935). Western Atlantic: Littorina ziczac: Lebour (1945); Abbott (1954); Lewis (1960); Marcus and Marcus (1963); Borkowski (1969). L. flava: Mar- cus and Marcus (1963). L. lineata: Borkowski (1969). L. lineolata: Borkowski (1969). L. melea- gris: Lewis (1960). Nodilittorina tuberculata: Le- bour (1945); Abbott (1954); Lewis (1960). Tecta- rius muricatus: Lebour (1945); Abbott (1954); Lewis (1960). Explanation to plate 325 (opposite page) Figs. 1-4. Littorina (Littorinopsis) scabra scabra (Linné), showing some of the variation in color and form exhibited by the shell of this species. 1. from Pulau Hantu, SW of Keppel Harbor, Singapore (USNM 660841); 2. from Can- daranan Id., Balabac, Philippines (USNM 233258); 3. from N. Queensland, Australia (USNM 149898); 4. from Bohaydulong Id., North Borneo (USNM 658031). Fig. 5. Littorina (Littorinopsis) carinifera (Menke) from Bombay, India (USNM 90470). Figs. 6, 7. Littorina brevicula (Philippi) from Awaji, Japan (USNM 343538). Figs. 8-10. Littorina (Littoraria) undulata Gray. 8. from Pulau Nias, SW Sumatra, Indonesia (USNM 654451); 9, 10. from N. shore of Guadalcanal Id., Solomon Ids. (USNM 598188). Figs. 11,12. Littorina (Littoraria) coccinea (Gmelin) from SW Viti Levu, Fiji (USNM 531795). Figs. 13,14. Littorina (Littoraria) mauritiana (Lamarck) from Mauritius (USNM 26744), Figs. 15,16. Littorina (Littoraria) pintado pintado (Wood) from Hilo, Hawaii (USNM 339401). Figs. 17,18. Littorina (Austrolittorina) unifasciata unifas- ciata Gray. 17. from Port Jackson, New South Wales (USNM 89472): 18. from Kalbarri, Western Australia (USNM 691672). Figs. 19,20. Nodilittorina australis (Gray). 19. from Rott- nest Id., Western Australia (USNM 671214): 20. from Port Denison, Western Australia (USNM 691677). Figs. 21,22. Nodilittorina nodosa (Gray). 21. from “Western Australia” (USNM 679494): 22. from Kalbarri, Western Australia (USNM 691680). Fig. 23. Littorina (Littoraria) kraussi Rosewater, from Mau- ritius (USNM 89410). Figs. 24,25. Nodilittorina pyramidalis (Quoy and Gaimard) from Lord Howe Id., off New South Wales, Australia (USNM 684715). Figs. 26,27. Littorina (Littoraria) praetermissa May from Port Fairy, Victoria, Australia (USNM 637353). Figs. 28,29. Littorina (Littorinopsis) melanostoma Gray. 28. from Pulau Lumut, Port Swettenham, Malaysia (USNM 661028); 29. from Kranji, Singapore (USNM 631930). [05-263] 420 Littorinidae B. Egg Mass— North and Northeastern Pacific: Littorina at- kana: Kojima (1958a, d); Habe (1958). L. sitkana: Habe (1958); Yamamoto and Habe (1962). Western and Eastern Atlantic: Littorina ob- tusata: Thorson (1946). C. Ovoviviparous— Indo-Pacific: Littorina scabra scabra: Whipple (1965); Struhsaker (1966). Eastern and Western Atlantic: L. scabra angu- lifera: Lebour (1945); Lenderking (1954); Marcus and Marcus (1963). L. saxatilis: Thorson (1946). Joseph Rosewater Littorinidae In general, based on available information, most members of the subgenera Littorina s.s. and Littoraria produce a plano-convex or bi-convex egg capsule, containing from one to a dozen or so eggs (L. littorea, brevicula, squalida, undulata, coccinea, pintado). Information regarding the na- ture of the capsule is lacking for some species, such as L. planaxis. In the cases of L. obtusata, sitkana and atkana an egg mass is deposited on the substrate and, of course, L. saxatilis is ovovi- viparous. Members of the subgenus Littorinopsis for which data are available are also ovovivipa- rous. In Austrolittorina and Nodilittorina rather Plate 326. Type-species of Genera included in Indo-Pacific Littorinidae, Part I. Figs. 1,2. Littorina (Littorina) littorea (Linné, 1758); speci- men figured in J. G. Jeffreys, 1865, British Conchology, Vol. 3, pl. 8, fig 3 (Jeffreys Collection, USNM 185523; 29.7 x 21.3 mm.) Figs. 3,4. Littorina (Littorinopsis) angulifera (Lamarck, 1822) from Sabanilla, Colombia (USNM 103151; 34.1 x 19.1 mm.) Fig. 5. Littorina (Austrolittorina) unifasciata unifasciata (Gray, 1826) from Port Jackson, Australia (USNM 89472; 20.9 x 12 mm.; for abapertural view see pl. 359, fig. 2). Figs. 6,7. Littorina (Littoraria) zebra (Donovan, 1825) from Venado River, Panama Canal Zone (USNM 589696; 29.6 x 19.9 mm.) Figs. 8,13. Nodilittorina (Granulilittorina) millegrana (Phi- lippi, 1848) from Pulau Jerak, West of Sembilan Islands, Malaysia (USNM 661049; 12.1 x 8 mm.). Figs. 9,11. Nodilittorina (Nodilittorina) pyramidalis (Quoy and Gaimard, 1832) from Airport Beach, Barrow Island, Western Australia (USNM 691681; 11.9 x 7.2 mm.) Figs. 10,12. Nodilittorina (Echinolittorina) tuberculata (Menke, 1828) from Permé, northwest of Cape Tiburon, Atlantic coast of Panama (USNM 664217; 16.4 x 11 mm.) [included for comparison, although there are no Indo- Specific species]. [05-264] November 30, 1970 elaborate ““gear-decorated” or “‘terraced” capsules are produced. Of the three species whose egg capsules are known in Melarhaphe, one produces a flattened cylinder-shaped capsule (L.neritoides) another a tiered capsule (L. meleagris) and the third a biconvex capsule (L. flava) [the generic assignments of the last two forms is still question- able]. Tectarius muricatus produces a capsule which is rather like that of L. neritoides, but is biconvex. A definite systematic trend in the shapes of littorinid egg capsules does not seem to exist Plate 327. Littorina littorea (Linné) removed from shell show- ing right side of body of A. female; B. male (from Fretter and Graham, 1962). Abbre viations— INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorininae 421 ag, albumen gland OV, ovary em, columellar muscle p, penis cov, covering gland png, penial glands cpg, capsule gland pr, prostate dg, digestive gland ro, renal section of oviduct e, eye sf, sole of foot ie foot sgr, sperm groove hg, hypobranchial gland © sn, snout k, kidney sts stomach me, mantle edge ts tentacle op, operculum td, testicular duct acting as opr, ovipositor vesicula seminalis above the species level. It does appear, however, that each species has fairly characteristic spawn. Radulae The radulae of Littorinidae may be described as generalized taenioglossate (see pl. 328). So far as is known most species browse on the epiphytes of the hard substrates upon which they live. The radulae are adapted for scraping off this food and may become very long as considerable replace- ment of worn teeth is required. The unused portion is carried coiled up in the radula sac behind the head. Radulae of Littorina and its subgenera are all fairly similar, having the formu- la 2-1-1-1-2, and consisting of a slender multicus- pid outer marginal, (pl. 328 D) a more robust inner marginal (pl. 328 C), a subquadrate lateral with a basal embayment (the embayment is char- acteristic of Littorinidae ) (pl. 328B), and a rachi- dian or central which is typically about as high as it is wide and which bears 3 cusps. In Nodilit- torina and Tectarius there is a tendency for narrowing of the central tooth although the lateral and marginals are quite similar to the condition in Littorina. In the genus Echininus and especially its subgenus Tectininus, as shown by Abbott (1954), there is considerable reduction in width of both central and lateral teeth. Since most mem- bers of the Littorinidae apparently share the gen- eralized taenioglossate radula and squarish lateral teeth, it does not seem to offer much promise for systematic diagnosis below the generic level. Distribution Most Littorinidae inhabit tropical seas, and the largest numbers of species live in the Indo- Pacific region. A conservative estimate places the total number of Recent world-wide Littorinidae of all genera at slightly above 100 species, exclud- ing the freshwater representatives which at least may be considered a separate subfamily of the Littorinacea as may the members of the Rissel- (UA) PAA? 7 7 F TO Da ee a os L_Olmm__ 5 Plate 328. Radula of Littorina littorea (Linneé), type-species of the genus Littorina. One complete transverse row of many which make up radula ribbon. Fig. A. rachidian or central tooth; Fig. B. lateral tooth C. and D. inner and outer marginal teeth. [05-265 ] 422 Littorinidae Joseph Rosewater Littorinidae lidae. Of these, some 34 species are to be found in the tropical Indo-Pacific, 21 inhabit the eastern Pacific, 16 the western Atlantic, only 9 the eastern Atlantic (5 of which are shared with the western Atlantic) and 27 more or less have been enumerat- ed in the South Ocean (Antarctic) (see List of Recognized Taxa). Representatives of the family are conspicuously absent from the southern tip of South America. The distributional pattern is further complicated by the present in two or more regions of what are termed analogues, species which bear considerable resemblance and may be related (Rosewater, 1963b). Since the geologic history of the Littorinidae dates from the Jurassic, it is doubtful that origins of the family can be unraveled. Dispersal of species is aided now and undoubtedly was helped in the past by pelagic larvae and the semi-arboreal habitat of such species as L. scabra. A single fertilized female of the latter oviparous species, if rafted to a favor- able locality, could establish a new colony. Classification Adapted as they are for intertidal life, it is not surprising to find little variation in general pat- tern of littorinid speciation. The shells are usually rounded-turbinate or some modification of this shape, permitting the animals to cling to rocks in surf or creep into crevices to avoid dessication. Grossly, the animals which form the shells do not vary much either. The systematists’ task of classi- fying species and groups of species into genera has come to depend on consistent differences in shape and sculpture and outstanding differences in anatomical details. Characters which have been utilized most in the present study are ab- sence or presence of nodulosity and granulate sculpture, overall color pattern and shape, the radula, and penile anatomy. The latter character appears to be an extremely important and reliable one, and two levels of classification have been based on it, at least in part: 1) the separation of species and 2) the formation of species groups (genera) exhibiting overall similarity in penile anatomy and other features. Difficulties have been encountered in the clas- sification of fossils. As mentioned previously, shells of Littorinidae resemble those of a number of other marine, freshwater and land _proso- branchs. If fossil preservation is not good, the decision on familial placement is extremely dif- ficult, if not impossible. Even if preservation is optimum, since there are no anatomical charac- ters upon which to rely, one can almost never be sure that these assignments are correct. This is especially true of the older fossils where resem- blance to Littorinidae is rather nebulous. For these reasons much concerning fossils has been taken on faith and must be regarded in that light. Only Tertiary Indo-Pacific fossils are included here or listed in the List of Recognized Taxa. As is the case with many groups of animals, not alone mollusks, an ample number of species and genera have been described in the Littorinidae. In many cases these are not justified in the light of modern systematics and some have had to be synonymized. Early in the course of this study it was thought probable that most of the generic names erected for species originally described or early assigned to Littorina would be of little value. After long evaluation, it has become ap- parent that some of these names may have real utility, at least as subgeneric groups indicators and it may be noted that they are so used through- out. It is hoped that in each case the reason for their use will be made clear. List of Recognized Taxa The following is a list of recognized species, subgenera and genera of world Littorinidae be- longing to the subfamily Littorininae. Fossils listed are Tertiary forms described from the Indo- Pacific region alone; the sheer numbers and the uncertainties surrounding extraterritorial and pre-Tertiary fossil species preclude their con- sideration here. As mentioned previously, the generic groupings are controversial but are of- fered as a working solution to some method of orderly arrangement of species. Due to the means of presentation many possibly familiar names will be missed by some in the list. Often they are considered to be synonyms and in the cases of Indo-Pacific species are included in the synony- mies which follow. The few fossil species names are preceded by a dagger [f]. Certain littorinoid groups, although definitely “Indo-Pacific” in general geography, are not con- sidered here. These are the members of such genera as Bembicium and Peasiella here believed to constitute at least a separate subfamily of Littorinidae. I consider the freshwater forms, such as Cremnoconchus and Cremnobates, also to be separable as at least a subfamily. Forms having definite southern ocean affinities, inhabiting New Zealand and the Antarctic, such as Laevilitorina, Macquariella, etc., are included in the list of recognized taxa, but will not be considered in detail here. A second part of Indo-Pacific Lit- torinidae, covering the subfamilies Tectariinae and Echininae will appear at some future time. [05-266] November 30, 1970 Family Littorinidae Gray, 1840 Subfamily Littorininae GENUS Littorina Ferussac, 1822 Subgenus Littorina Ferussac, 1822. littorea (Linné, 1758). Type. Recent, North Atlantic obtusata (Linné, 1758). Recent, North Atlan- tic saxatilis (Olivi, 1792). Recent, circumboreal sitkana Philippi, 1846. Recent, northeast Pacific atkana Dall, 1886. Recent, North Pacific aleutica Dall, 1872. Recent, North Pacific squalida Broderip & Sowerby, 1829. Recent, North Pacific brevicula (Philippi, 1844). Recent; East Asia, Japan planaxis Philippi, 1847, Recent, northeast Pacific scutulata Gould, 1849. Recent, northeast Pacific ziczac (Gmelin, 1791). Recent; tropical wes- tern Atlantic Subgenus Littoraria Gray, 1834 zebra (Donovan, 1825). Type. Recent, tropi- cal eastern Pacific undulata Gray, 1839. Recent, Indo-Pacific nebulosa (Lamarck, 1822). Recent, Carib- bean and Gulf of Mexico mauritiana (Lamarck, 1822). Recent, Indian Ocean kraussi Rosewater, new name Recent, Indian Ocean coccinea (Gmelin, 1791) Recent; Indo- Pacific praetermissa May, 1909. Recent, South Aus- tralia pintado (Wood, 1828). Recent; Indo-Pacific pintado schmitti, Bartsch & Rehder, 1939. Recent, Clipperton Island pullata Carpenter, 1864. Recent; tropical eastern Pacific cingulifera Dunker 1845. Recent; tropical west Africa ?sundaica Altena, 1945. Recent, Java Pacutispira E. A. Smith, 1892. Recent, Aus- tralia Pinfans E. A. Smith, 1892. Recent, Australia tkozaiensis Nomura and Onisi, 1940. Mi- ocene, Japan tadonis Yokoyama, 1927. Pliocene, Japan tlucida Yokoyama, 1927. Pliocene, Japan INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorininae 423 Subgenus Littorinopsis Mérch, 1876 scabra scabra (Linne, 1758). Recent, Indo- Pacific scabra angulifera (Lamarck, 1822). Type. Recent, eastern and western Atlantic scabra aberrans Philippi, 1846. Recent, trop- ical east Pacific melanostoma Gray, 1839. Recent, south Asia. carinifera (Menke, 1830). Recent, Indo- Pacific irrorata (Say, 1822). Recent, western Atlan- tic fasciata Gray, 1839. Recent, tropical east Pacific varia Sowerby, 1832. Recent, tropical east Pacific modesta Philippi, 1846. Recent, tropical east Pacific paytensis Philippi, 1847. Recent, tropical east Pacific + miodelicatula Oyama, 1950. Miocene, Ja- pan. tincisa Yokoyama, 1927. Pliocene, Japan. Subgenus Austrolittorina Rosewater, new sub- genus unifasciata unifasciata Gray, 1826. Type. Recent, Australia unifasciata antipoda Philippi, 1847. Recent, New Zealand unifasciata fernandezensis Rosewater new subspecies. Recent, Islas Juan Fernandez cincta Quoy and Gaimard, 1832. Recent, New Zealand araucana Orbigny, 1840. Recent, Chili and Peru peruviana (Lamarck, 1822). Recent, south- eastern Pacific aspera Philippi, 1846. Recent, tropical east Pacific penicillata Carpenter, 1864. Recent, Baja California tessellata Philippi, 1847. Recent, Caribbean lineata Orbigny, 1841. Recent, tropical wes- tern Atlantic lineolata Orbigny, 1840. Recent, tropical western Atlantic punctata (Gmelin, 1791). Recent, East At- lantic and South Africa africana Philippi, 1847. Recent, South Afri- ca knysnaensis Philippi, 1847. Recent, South Africa 424 Littorinidae Subgenus Melarhaphe Menke, 1828 neritoides (Linne, 1758) Type. Recent, east Atlantic ?mespillum (Muhlfeld, 1824). Recent, tropi- cal western Atlantic ?meleagris (Potiez and Michaud, 1838) Re- cent, West Indies ?umbilicata Orbigny, 1840. Recent, Chili and Peru ? flava King and Broderip, 1832. Recent, tropi- cal west Atlantic Subgenus Algamorda Dall, 1918 newcombiana (Hemphill, 1877). Type. Re- cent, northwest North America GENUS Nodilittorina von Martens, 1897 Subgenus Nodilittorina von Martens, 1897 pyramidalis pyramidalis (Quoy and Gai- mard, 1833). Type. Recent, Indo-Pacific. pyramidalis pascua Rosewater, new subspe- cies. Recent, Easter and Pitcairn Islands. natalensis (Philippi, 1847). Recent, East Af- rica and Madagascar. australis (Gray, 1826). Recent, W. Australia. nodosa (Gray, 1839). Recent W. Australia. galapagiensis (Stearns, 1892). Recent, Gala- pagos Islands. Subgenus Echinolittorina Habe, 1956 tuberculata (Menke, 1828). Type. Recent, tropical west Atlantic. Subgenus Granulilittorina Habe and Kosuge, 1966 millegrana (Philippi, 1948). Type. Recent, Indo-Pacific. subnodosa (Philippi, 1847). Recent, Red Sea and Persian Gulf. leucosticta leucosticta (Philippi, 1847). Re- cent, India. leucosticta biangulata (von Martens, 1897). Recent, east Indies and western Melane- sia. leucosticta feejeensis (Reeve, 1857). Recent, western Pacific. exigua (Dunker, 1860). Recent, Japan. tiwakiana (Nomura and Hatai, 1936). Mi- ocene, Japan. cinerea (Pease, 1869). Recent, Marquesas Islands picta (Philippi, 1846). Recent, Hawaii. Joseph Rosewater Littorinidae ?miliaris (Quoy and Gaimard, 1833). Recent, east Atlantic ?striata (King and Broderip, 1832). Recent, east Atlantic GENUS Laevilitorina Pfeffer, 1886 Subgenus Laevilitorina Pfeffer, 1886 caliginosa caliginosa (Gould, 1849). Type. Recent, Southern Ocean caliginosa aestualis Strebel, 1908. Recent, Antarctica antipoda (Filhol, 1880) Recent, New Zea- land bifasciata Suter, 1913. Recent, New Zealand bennetti Preston, 1912. Recent, Antarctic claviformis Preston, 1916. Recent, Antarctic granum Pfeffer in von Martens & Pfefter, 1886. Recent, Antarctic latior Preston, 1912. Recent, Antarctic pygmaea Pfefter, 1886. Recent, Antarctic umbilicata Pfeffer, 1886. Recent, Antarctic venusta Pfeffer, 1886. Recent, Antarctic antarctica E. A. Smith, 1902. Recent, Ant- arctic Subgenus Corneolitorina Powell, 1951 coriacea (Melvill and Standen, 1907). Type. Recent, Antarctic elongata Pelseneer, 1903. Recent, Antarctic heardensis Dell, 1964. Recent, Antarctic GENUS Macquariella Finlay, 1926 hamiltoni (Smith, 1898). Type. Recent, New Zealand aucklandica Powell, Zealand delli Powell, 1955. Recent, New Zealand macphersonae Dell, 1964. Recent, Mac- quarie Island 1933. Recent, New GENUS Laevilacunaria Powell, 1951 Subgenus Laevilacunaria Powell, 1951 bransfieldensis (Preston, 1916). Type. Re- cent, Antarctic antarctica (von Martens, 1885). Recent, Ant- arctic pumilio (E. A. Smith, 1879). Recent, Antarc- tic Subgenus Pellilacunella Powell, 1951 bennetti (Preston, 1916). Type. Recent, Ant- arctic [05-268] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorininae 425 GENUS Pellilitorina Pfeffer, 1886 setosa (Smith, 1875). Type. Recent, Antarc- tic pellita (von Martens, 1885) Recent, Antarc- tic GENUS Rissolittorina Ponder, 1966 alta (Powell, 1940). Type. Recent, New Zea- land mariae (Tenison-Woods, 1876). Recent, Tas- mania Abbreviations The following institutional abbreviations are used in this paper: AMS-Australian Museum, Sydney ANSP-Academy of Natural Sciences of Phil- adelphia BM(NH)-British Museum (Natural History}, Lon- don BPBM-Bernice P. Bishop Museum, Honolulu MHNG-Muséum d Histoire Naturelle, Geneva MHNP-Muséum d'Histoire Naturelle, Paris MCZ-Museum of Comparative Zodlogy, Cam- bridge, Massachusetts NMW-National Museum of Wales, Cardiff RNHL-Rijksmuseum van Natuurlijke Historie, Leiden SMF-Senckenberg Museum, Frankfurt-am-Main USNM-United States National Museum, Wash- ington, D.C. WAM-Western Australian Museum, Perth, West- ern Australia YPM-Peabody Museum of Natural History, Yale University, New Haven, Connecticut ZMA-Zoological Museum, Amsterdam ZMC-Zoological Museum, Copenhagen Acknowledgements The following persons contributed to making this study more complete by providing informa- tion, research assistance, specimens or working space; their help is gratefully acknowledged: R. T. Abbott, Delaware Museum of Natural History; C. O. van Regteren Altena, E. Git- tenberger, R. M. van Urk-RNHL; E. Binder- MHNG; J. B. Burch-Museum of Zoology, Uni- versity of Michigan; W. J. Clench, R. D. Turner, kK. J. Boss-MCZ; H. Coomans-ZMaA; S. P. Dance- NMW; R. K. Dell-Dominion Museum, Wel- lington, New Zealand; E. Fisher-MHNP; L. Fish- elson-Tel-Aviv University, Israel; W. D. Hart- man-YPM; Johnny Johnson and Mr. and Mrs. D. Everett-Singapore; A. M. Keen-Stanford Univers- ity; J. Knudsen, H. Lemche-ZMC; Y. Kondo- BPBM; S. Kosuge-National Science Museum, Tokyo; H. S. Ladd-U. S. Geological Survey; D. F. McMichael-AMS; R. Robertson and V. Orr Maes- ANSP; H. Steinitz-Hebrew University, Jerusa- lem, Israel; Jeannette Whipple Struhsaker- University of Hawaii, Honolulu; N. Tebble, J. F. Peake-BM(NH); G. Thorson-Marine Biological Lab, Helsinggr, Denmark; B. R. Wilson-WAM; A. Zilch-SMF; Mrs. Ann Cohen called to my atten- tion specimens she collected in the Juan Fer- nandez Islands; E. Brody, A. Griffin, J. Green- field, R. Capen, Research Assistants; H. A. Rehder, J. P. E. Morrison, C. F. E. Roper, Divi- sion of Mollusks, USNM. The U.S. Atomic Energy Commission, through I. E. Wallen and R. W. Hiatt, arranged for me to work at the Eniwetok Marine Biological Labora- tory. A portion of this work was supported by the National Science Foundation as a part of the U.S. Program in Biology, International Indian Ocean Expedition. The work was also supported in part by Smithsonian Institution Research Foundation Grant number Sg 0684025. Doubtful Species of Littorinidae Included under this heading are species of Littorinidae from the Indo-Pacific which I con- sider either unrecognizable as valid species from the faunal area, or as being questionably included in the family. In the latter category, some species already have been removed from the family by prior workers. Although Iredale and McMichael (1962) listed Larinopsis turbinata (Gatliff and Gabriel, 1909) and L. ostensus Iredale, 1936, in Littorinidae, the genus Larinopsis Gatliff and Gabriel, 1916, was placed in Fossaridae by Wenz (1940) and by Macpherson and Gabriel (1962, including the mentioned species). Ponder (1966) has shown that some species placed in Zelaxitas, formerly thought to be Littorinidae, belong to three different families, that genus now being assigned to Rissoellidae. Only one species origi- nally included under Zelaxitas, Z. alta Powell, 1940, remains in Littorinidae, although in a dif- ferent genus (see List of Recognized Taxa under Rissolittorina Ponder). Littorina beccarii Tapparone-Canefri, 1875 Range—Known only from the type-locality: So- rong, northwest New Guinea (West Irian, Indone- sia); [05-269] 426 Littorinidae Remarks—This species has not been figured and its identity is in doubt. The author’s sugges- tion that an affinity exists with Littorina lamellosa Montrouzier indicates that it is a Fossarus and not a Littorina. Mrs. van Benthem Jutting (1962) published the results of her studies on the types of New Guinea non-marine mollusks described by Tapparone-Canefri. These were preserved in the Genoa Museum, and it is possible that the type of L. beccarii is there also. Unfortunately it has not been possible to ascertain this fact in the course of the present study Synonymy— 1875 Littorina beccarii Tapparone-Canefri, Annali del Museo civico di Storia Naturale di Genova, vol. 7, p. 1031 (Sorong [N.W. New Guinea]). Littorina fragilis Fenaux, 1943 Remarks—The illustration accompanying the description of this species indicates that it is very probably a member of the genus Diala (Ce- rithiidae) and not a Littorina. Synonymy— 1943 Littorina fragilis Fenaux, Bulletin de l'Institut océan- ographique, Monaco, no. 835, p. 7, figs. 4,5 (Pau- motou). Problitora globulus (Angas, 1880) Remarks—Macpherson and Gabriel (1962) placed this species in the genus Problitora Ire- dale, 1931, of which P. moerchi (Adams and Angas) is the type-species. As globulus was de- scribed from shell-sand, and has been collected on only very few occasions, its true relationships remain to be proven. At present there appears to be little reason for retaining these peculiar forms in the family Littorinidae as they resemble much more closely members of the family Naticidae. Synonymy— 1880 Amauropsis globulus Angas, Proceedings of the Zoologi- eal Society of London, 1880, part 3, p. 416, pl. 40, fig. 5 (Holdfast Bay in shell-sand [near Adelaide, South Australia]); types, BM(NH) 81.4.29.5. Littorina hisseyiana Tenison-Woods, 1876 Remarks—May (1903) figured the type of L. hisseyiana. His illustration shows a tiny, ap- parently umbilicated snail with a multispiral operculum, and having a color pattern resembling Joseph Rosewater Littorinidae one of the Trochacea. It is certainly not a Lit- torina. Synonymy— 1876 Littorina hisseyiana Tenison-Woods, Papers and Pro- ceedings of the Royal Society of Tasmania, for 1875, p. 148 (from stomach of a mullet, Agenostoma die- menensis Rich, probably caught in the Derwent [Tas- mania]). 1903, W. L. May, ibid., for 1902, p. 111, fig. 7; 7 syntypes in Tasmanian Museum (T.M. 5480) according to Hardy, 1916, ibid., for 1915, p. 66. Littorina lamellosa Montrouzier, 1861 Remarks—Although described as a Littorina, the author observed that this species resembles a fossarid. This is certainly the case, as the original illustration almost exactly matches the Hawaiian species which was named by Pease as Fossarus garretti. Synonymy— 1861 Littorina lamellosa Montrouzier, Journal de Conchyliol- ogie, vol. 9, p. 273, pl. 11, fig. 5 ({Ile] Art, New Caledonia). Problitora moerchi (Adams and Angas, 1864) Remarks—See Remarks under P. globulus (An- gas). I consider this species not to belong in Littorinidae. The figures by Hedley (1902) es- pecially that of the operculum, resemble more closely one of the Naticidae. Synonymy— 1864 Amauropsis moerchi Adams and Angas, Proceedings of the Zoological Society of London, 1863, part 3, p. 423 (Watson’s Bay, Port Jackson [New South Wales]); type, BM(NH) 70.10.26.178. 1902, Hedley, Proceed- ings of the Linnean Society of New South Wales, vol. 26, part 4, p. 700, pl. 34, figs. 19, 20. Littorina reticulata Anton, 1839 Remarks—Anton’s species could be any of sev- eral Granulilittorina. The description and Philip- pis figure are too vague to identify the species with any precision and the lack of a locality increases the likelihood that L. reticulata may be an extra-Indo-Pacific entity. The species is here considered unrecognizable. Synonymy— 1839 Litorina reticulata Anton, Verzeichniss der Conchylien welche sich in der Sammlung von H. E. Anton befinden. Halle, p. 53 (no locality given). 1847, Phi- lippi, Abbildungen und Beschreibungen Conchylien, vol. 2, p. 199, pl. 4, fig. 12 [figure said to be drawn from one of Anton’s type-specimens]. 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Publications of the Seto Marine Biological Laboratory, vol. 1, no. 3, pp. 151-152, fig. Tokioka, T. and T. Habe. 1953. A new type of Littorina- capsula. ibid., vol. 3, no. 1, pp. 55, 56, fig. Tryon, G. W. 1887. Family Littorinidae. Manual of Concholo- gy, vol. 9, pp. 229-313, pls. 40-51. Von Martens, E. 1887. List of the shells of Mergui and its Archipelago, collected for the Trustees of the Indian Muse- um, Calcutta by Dr. John Anderson, F.R.S., Superintendent of the Museum. The Journal of the Linnean Society, Zo- ology, vol. 26, no. 130, pp. 155-219, 3 pls. Weinkauff, H. C. 1856-1882. Die Gattung Litorina. Systemat- isches Conchylien-Cabinet, edition 2, vol. 2, prt. 9, pp. 1-114, 14 pls. Weinkauff, H. C. 1883. Catalog der Gattung Litorina Feru- ssac. Jahrbuch der Deutschen Malakozoologischen Gesell- schaft, vol. 10, pp. 213-227. Wenz, W. 1940. Handbuch der palaozoologie, vol. 6, prt. 1 (6), pp. 721-960, figs. 2084-2787. Whipple, Jeanette A. 1965. Systematics of the Hawaiian Littorina Férussac (Mollusca: Gastropoda). The Veliger, vol. 7, no. 3, pp. 155-166, pls. 25, 26, 4 text figs. Whipple, J. A. [is maiden name of J. W. Struhsaker, q.v.]. Winckworth, R. 1922. Nomenclature of British Littorinidae. Proceedings of the Malacological Society of London, vol. 15, prts. 2, 3, pp. 95-97. Yamaguchi, M. 1967. Egg capsules of a Periwinkle, Littorina brevicula, in plankton samples. The Venus, vol. 25, no. 2, pp. 73-76, figs. 1, 2. Yamamoto, G. and T. Habe. 1962. Fauna of shell-bearing mollusks in Mutsu Bay. Scaphopoda and Gastropoda (1). Bulletin of the Marine Biological Station of Asamushi, Tohoku University, vol. 11, no. 1, pp. 1-20, pls. 1-3. Yen, Teng-Chien. 1942. A review of Chinese Gastropods in the British Museum. Proceedings of the Malacological Society of London, vol. 24, parts 5, 6, pp. 170-289, pls. 11-28. [05-277] 430 Littorinidae Joseph Rosewater Littorinidae Subfamily Littorininae Gray, 1840 Key To Subgeneric Groups of Indo-Pacific Littorininae The following key is an attempt to provide a point of departure for the placement of a littorinid species in a particular subgeneric group. Even at the generic level there is a great deal of phe- notypic variation expressed and it is often true that such hopefully dependable characters as nodular and granular sculpture are evanescent. Other keys are provided in the appropriate places to species of Littoraria and Littorinopsis. None are provided for the other groups, however, be- cause it was felt that species of Austrolittorina and Nodilittorina s.s. are sufficiently distinct and geographically situated so as to essentially pre- clude difficulties in identification. In the case of Granulilittorina, the nature of speciation in the group makes the construction of an effective key a near impossibility. It is suggested that the user may wish to acquire some familiarity with the range of variation in this subfamily before pro- ceeding. 1. Shell nodulose, granulose, or with surface axially folded ......... 0.00.00 ceca 2 1. Shell surface smooth or spirally folded (carinate) .. 0.0.0.0... 00 eee ee ee ee 2. Sculpture nodulose, limited to 2-3 rows per whorl of major nodules, or surface axially folded ee ee ee ee Nodilittorina p. 05-375 2. Sculpture granulose, often in excess of 3 rows of granulations, sometimes nearly smooth lite Bn BuO oniak te a eg 4A ace, BAe Op ats Be ies Soles Granulilittorina p. 05-395 3. Base of shell adjacent to columella flattened or hollowed out, forming a crescent-shaped BIS co deashs Cale Bee oe SE Rae Sa eee ea ee Ae Owes Austrolittorina p. 05-351 3. Base of shell adjacent to columella generally smoothly rounded, not flattened OO SC ONDE - seid hh a ae Ge eek Gods bee bs oe ea OEE BESS SRB OSu Cte ESOG 4 4. Spire less than half the length of shell, multiple penial glands present ahed Bad Se Sea Gee Bee ws dG ee a oa wee wees Littorina p. 05-285 4, Spire usually more than half the length of shell, penial glands absent ...........5 5. Shell rather thin for its size, conspicuously colored or patterned, usually arboreal. or plami-living Species: fei. avulk eee be Dene BS Littorinopsis p. 05-329 5. Shell not thin for its size, often with subdued coloration or lacking pattern, usually roclk-livine Species’ 24 dcn cdo beee nd deena ee oS ea Littoraria p. 05-297 November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 431 Family Littorinidae Gray, 1840 Subfamily Littorininae Gray, 1840 Genus Littorina Férussac, 1822 Type: Littorina littorea (Linneé, 1758) Considered broadly, the genus Littorinaencom- passes a multitude of Recent and fossil species throughout the world. The impossibility of prop- erly classifying the large number of poorly pre- served fossil species described from the Upper Paleozoic through the Tertiary prevents coverage here of any but Tertiary Indo-Pacific fossils. Living species of the genus Littorina usually occupy habitats on rocks in the intertidal zone although some species prefer shore vegetation. The geographic range of the genus is from Arctic to Antarctic shores, including intervening tem- perate and tropical areas where a suitable habitat is available. The subgenus Littorina, sensu stricto, however, is a northern group and there is only one species of that subgenus whose range extends into the tropical Indo-Pacific, Littorina brevicula (Philippi), which is treated here (see List of Recognized Taxa). Synonymy— 1822 Littorina Ferussac, Tableaux Systematiques des Ani- maux Mollusques, p. xxxiv. Type-species by subse- quent designation, Blainville, 1828, Dictionnaire des Sciences Naturelles, Paris, vol. 56, p. 98: Turbo littoreus [= Littorina littorea (Linne)]. 1827 Littorina “Ferussac’ Nilsson, Petrificata Suecana Forma- tionis Cretaceae, part 2, Mollusca, p. 11. Invalid type-species by monotypy: Turbo sulcatus Nilsson, 1827, a species not originally included by Feérussac. 1827 Neritoides T. Brown, Illustrations of the Conchology of Great Britain and Ireland, Index p. iv, pl. 43, figs. 14, 15, 21, 22. Type-species by monotypy: Neritoides littoralis [=Littorina obtusata (Linne, 1758)]. 1828 Litorina Menke, Synopsis Methodica Molluscorum, p. 24 [emendation of Littorina Ferussac, 1822]. 1847 Littorelaea Leach, Annals and Magazine of Natural History, vol. 20, p. 271; in combination Littorelaea pultneyii [nomen nudum|. 1849 Litormia Morch, Indbydelsesskrift til de oftentlige af- gangs-og aarsprgver i Nykjdbing katedralskole, p. 63 {error for Littorina Feérussac, 1822]. 1854 Bacalia H. and A. Adams, Genera of Recent and Fossil Mollusks, vol. 1, p.312; based on Bacalia Gray, 1840, Synopsis of the Contents of the British Museum, ed. 42, p. 147 [nomen nudum]; type-species by subse- quent designation, Winckworth, 1922: Turbo lit- toreus Linne. 1861 Littorrina Gabb, Proceedings of the American Philo- sophical Society, vol. 8, p. 114 [error for Littorina Férussac, 1822 1869 Neritrema Récluz, Actes de la Société Linnéenne de Bordeaux, series 3, vol. 7 (vol. 27) pp. 43, 46. Type- species by subsequent designation, Dall, 1909, U.S. Geological Survey Professional Paper 59, p. 79: Li- ttorina obtusata Linne,. 1878 Litonia Weinkauff, Systematisches Conchylien-Cabinet, vol. 2, part 9, p. 40 [error for Littorina Ferussac, 1822] 1884 Litorinna Dall, Proceedings U.S. National Museum, vol. 7, p. 344 [error for Littorina Ferussac, 1822] 1891 Isonema ‘Hall’ Provancher, Faune Canadienne; Les Mollusques de la Province de Quebec, part 1, p. 91. Type-species by subsequent designation Bequaert, 1943 Johnsonia, vol. 1, p. 1: Turbo littoreus Linne, 1758; not Isonema Meek and Worthern, 1866. 1891 Littornia Jenkins and Grocock, Hardwicke’s Science Gossip, vol. 27, p. 9; 1917, H. B. Preston, Zoological Record, vol. 52, section 8, Mollusca, p. 33 [error for Littorina Férussac, 1822]. 1918 Algaroda Dall, Proceedings of the Biological Society of Washington, vol. 31, p. 137; type-species by original designation: Littorina littorea Linne. 1918 Littorivaga Dall, ibid., Type-species by original designa- tion: Littorina sitchana Philippi. 1939 Neritotrema ‘Recluz’ Wenz, Handbuch der Palaozoo- logie, Lief 4, Band 6, Gastropoda, part 3, Proso- branchia p. 518 [emendation of Neritrema Récluz, 1869]. 1958 Exzolittorina Habe, Publications from the Akkeshi Ma- rine Biological Station, no. 8, p. 9; type-species by monotypy: Ezolittorina squalida Broderip and Sow- erby. As has been accurately pointed out by Winck- worth (1922) and Bequaert (1943) Férussac cited the names of several species in association with his description of Littorina. The “Paludina ma- rine’ listed on pages 9-11 of the introduction to Férussac’s work are obtusatus, neritoides, Lit- toreus, muricatus and afer. Following the list Férussac mentioned (p. 11) in vernacular “‘lit- torine’ as a subgenus to contain some species previously assigned to Turbo and Trochus. In the following paragraph he assigned Turbo neritoides of Linné to this subgenus, at least in part. Since the latter cannot be construed as a type designa- tion, it is necessary to seek a later one. Blainville (1824, Dictionnaire des Sciences Naturelles, Par- is, vol. 32, p. 226) gave as an example of Férus- sac’s “Littorine’: Turbo littoralis, referring to Chemnitz vol. 5, t. 185, fig. 1852, nos. 1-18 [1-8] [figures are L. littorea]. This is an invalid subse- quent designation (I.C.Z.N. Art. 69 (a) (i) and (iv)), and the same is true in the case of Pay- raudeau's (1826) description of L. basterotii [=L. neritoides (Linné)]| in association with Littorina. The first valid designation is that of Blainville, (1828 ibid.) who clearly indicated Turbo littoreus, one of the originally included species as the type of the genus. [05-279] 432 Littorinidae Joseph Rosewater [ These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence. | [05-280] Littorinidae November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 433 Subgenus Littorina sensu stricto Characteristics of species belonging to the sub- genus Littorina s.s. include a generally boreal distribution, relatively large, somber colored shells (with a number of exceptions) and the morphological character of the presence in the male of a series of penial glands with accessory flagella on the penis. Littorina brevicula (Philippi, 1844) (Pls. 325, 329, 330) Range—Northern China, Korea and Japan (and boreal Pacific). Remarks—Littorina (Littorina) brevicula is an inhabitant of the boreal Pacific. Although strictly a cool water species, it is included here with species occuring in the tropical Indo-Pacific be- cause its range does extend to the southern most of Japan’s main islands and also to the coast of China at latitudes roughly on a level with Tai- wan. It occurs therefore within the range of a number of tropical species. The anatomy of the penis of L. brevicula, which possesses multiple penial glands, the char- acteristics of the floating egg (helmet) capsule, which is a relatively simple plano-convex shape, the generalized radula and somber coloration of the shell all align this species with the boreal group of Littorina s.s. Habitat—Intertidal on rocky coasts. Plate 329. Littorina brevicula (Philippi). Figs. 1,2. Specimens from Takami, Honshu, Japan (USNM 424296; 11.7 x 9.3 mm.) Figs. 3,4. Littorina souverbiana Crosse, Holotype (Journal de Conchyliologie Collection, Paris Museum; 11.5 x 11 mm.) Figs. 5,6. Turbo (Litorina) breviculus Philippi, lectotype figure (from’ Abbildungen und Beschreibungen Con- chylien, vol. 2, p. 161, pl. 3, fig 10; about 11 x 10 mm.) Figs. 7,8. Littorina balteata Reeve, lectotype (British Muse- um (NH) 1968316; 11 x 7.6 mm.). Description—Shell reaching nearly 23 mm. (about 0.8 inch) in length, broadly-to depressed turbinate in shape, often appearing almost as broad as long, but having an average obesity of about .74 (39 specimens range from .68-.79): rather thick in structure, imperforate; usually displaying an accentuatedly pointed siphonal trough; body whorl often sculptured with 3 or 4 raised strong, white-speckled spiral cords. Ex- ternal color variable; reddish brown to grayish white solid color or with banding, spotting, strip- ing and zigzag color patterns. Aperture diffuse tan to brown with faint narrow white band revolving inward from near anterior junction of outer lip and columella, occasionally a wider, very diffuse band visible inside outer lip midway to its anteri- or junction with columella; columella grayish white anteriorly, merging with aperture color posteriorly; inner edge of outer lip often dark brown or speckled brown and white. Base dis- tinctly flattened with comparatively low spiral cords; whorls broadly shouldered below suture. Whorls 4-5, rounded, nuclear whorls usually e- roded. Spire usually less than half the length of shell, produced at an angle varying from about 75° to 110° in very shortspired individuals. Aper- ture rounded-oval; inner and outer lips relatively thickly produced; with a distinctly pointed si- phonal trough at anterior junction of outer lip and columella, often tinged with brown in young individuals; base adjacent to columella seldom depressed, but occasionally an anomalous umbil- ical chink formed. Suture impressed. Spiral sculpture usually consisting of from 3-4 major raised cords on upper half of body whorl; from Plate 330. Littorina brevicula (Philippi). Fig. A. Radula, showing central and one-half row of teeth (upper scale is 0.125 mm.); Hakodate, Japan. Fig. B. Penis of young specimen from Wuchiuhsu, China (lower scale is 1 mm.) [05-285] 434 Littorina Littorinidae Joseph Rosewater 4-6 minor cords on lower half on base; spire sculpture usually at least partially obliterated. Nuclear whorls about 2-2!/2, smooth, light- brown; post nuclear whorls rapidly becoming spirally sculptured with 1-2 cords. Overall micro- scopic sculpture consisting of fine wavy spiral threads especially noticeable between major cords, and fine, rather regular, oblique axial growth markings. Operculum roundly oval, pau- cispiral, the nucleus wen. Peas and nearly central. Radula littorinoid, 2-1-1-1-2, the central tooth broad. Animal darkly pigmented on surfaces of tenta- cles, snout and foot; the penis of small individu- als (shell length 5-6 mm.) bearing 2 or 3 penial glands with accessory flagella. Reproduction oviparous, producing pelagic egg capsules, plano-convex in shape. The capsule measured about 350m in maximum diameter and 170m in height, the egg about 84 in diameter, there being usually only 1 egg per capsule (Kojima, 1957). The breeding season was found to be from Febru- ary to April at Asamushi, Japan. Measurements (mm)— length width no. whorls locality 22.8 16.0 4 Hakodate, Japan 20.1 13.7 4+ Kamoito, Teshiwo, Japan 19.3 15,2 4 Muroran, Japan 17.8 13.3 4 Muroran, Japan 16.2 12.0 4 Awaji, Japan 14.4 11.4 4 Muroran, Japan 12.6 9.2 5 Shirahama, Japan 11.2 8.5 3 Side Saddle Island, China 10.8 8.5 4 Cheetfoo, China 9.3 6.8 4 Pei-tai-ho, China Synonymy— 1844 Turbo (Litorina) breviculus Philippi, Zeitschrift fur Malakolozoologie, vol. 1, p. 166 (China, at mouth of Yang-tse-Kiang (Largilliert!); 1847, Abbildungen und Beschreibungen Conchylien, vol. 2, p. 161, Litorina, pl. 3, fig 10. (lectotype figure). 1857 Littorina balteata Reeve, Conchologia Iconica, vol. 10, Littorina, pl. 14, fig 71, a,b (China; lectotype BM(NH) 1968316: 11 x 7.6 mm.). 1861 Litorina mandshurica Schrenck, Bulletin de L’Académie Impériale des Sciences de St. Pétersbourg, vol. 4, no. 7, p. 409 (Sinus des Castries [Zaliv Chikhacheva, Gulf of Tartary, USSR]) type in Academy of Sciences Leningrad? 1867, Reisen und Forschungen im Amur-Lande, vol. 2, p. 333, pl. 14, figs 14-20. 1862 Littorina souverbiana Crosse, Journal de Conchyliologie, vol. 10, p. 53, pl. 1, figs 6-7 (in Sinu Talienwhanensi [North China]; holotype MHNP: 13 x 10 mm), 1867 Litorina mandshurica carinata Schrenck, Reisen und Forschungen im Amur-Lande, vol. 2, p. 338 (no locality given). 1867 Litorina mandshurica laevigata Schrenck, ibid. 1887 Littorina manschurica ‘Schrenck’ Tryon, Manual of Conchology, vol. 9, pp. 241, 300 [error for L. mand- shurica Schrenck]. 1928 Littorina heterospiralis Grabau and King, Shells of Peitaiho, Peking Society of Natural History, Hand- book no. 2, second ed., p. 230, pl. 10, fig. 104 (Peitaiho [China]); types in Peking? Types—The type specimen of Turbo (Litorina) breviculus Philippi may be in the Berlin Muse- um, although it has been impossible for me to determine this with certainty. The lectotype fig- ure appears in Abbildungen und Beschreibungen Conchylien, pl. 3, fig 10 (see our pl. 329 figs. 5, 6). The lectotype of Littorina balteata Reeve is in the British Museum (NH)1968316. Types of L. mandshurica Schrenck and of its subspecies car- inata and laevigata, may be in the Academy of Sciences, Leningrad. The holotype of L. souver- biana Cross is in the Paris Museum (Journal de Conchyliologie Collection). The type of L. heter- ospiralis Grabau and King, may be in Peking. Records—KOREA: Wonsan Beach, Wonsan (MCZ); 2 mi. W. of Masan (USNM); Sorai (Sorai-men) (ANSP). CHINA: Port Arthur (Lu-shuh), Manchuria (ANSP, MCZ); Pei-tai-ho, N. Chihli coast; Cheefoo (Yen T’ai); Chiaochou (Kiautschou), nr. Tsingtau (Isington), Yellow Sea; Side Saddle Island (Lu- Hua Shan), Che-Kiang Province; Foochow (Fu-chou) (all USNM); Wuchiu Hsu (ZMC). JAPAN: HONSHU: Kominato; Yenosima (Islands) (USNM, MCZ); Isu, Ise (Isu-shima?) (MCZ):; Tokyo Bay (Yedo Bay) (ANSP, MCZ); Tsukishima, Tokyo, Tokyo Bay (USNM, MCZ); Shirahama, Wakayama Prefecturate; Yokohama (both ANSP, USNM); Misaki, Sagami Bay, off Hondo (USNM); Sagami Bay, Zushi Beach (MCZ); Shinagawa Bay, Tokyo; Toshima, Tanabe-wan, Seto Marine Biology Lab., Shirahama, Wakayama-ken (both USNM); In- land Sea (Seto-Naikai) (MCZ, ANSP); Awaji-shima (Island) (USNM, MCZ); Wakanura; Takami (both USNM). KYUSHU: Nagasaki; Camp Mower, Sasebo (both USNM); Hirado, Hizen (ANSP, USNM): Shimonoseki; Saigo, Dogo Islands, Oki Group, Japan Sea, shore (both USNM). HOKKAIDO: Muro- ran (USNM); Hakodate (USNM, MCZ); Zenibako (BPBM); Otaru; Kamoito, Teshio (both USNM). i aoe ae \4o0° 160° f i “ 4 PHILIBPINE “Guar of 7, a 3 Enwefok. > ce J) LU? | Oth Bhs Wg ee ‘ . . . > ws ee : CAROLINE ae te Plate 331. Geographical distribution of Littorina brevicula (Philippi.) sl [05-286] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 435 Subgenus Littoraria Griffith and Pidgeon, 1834 Type: Littorina zebra (Donovan, 1825) Members of this subgenus are inhabitants of tropical and subtropical portions of the Indo- Pacific, East Pacific and West Atlantic, but repre- sentatives appear to be absent from the East Atlantic. Although the type-species of Littoraria is from tropical west America, the majority of species live in the Indo-Pacific with one species in the Caribbean area. This may reflect what appears to be a trend toward a higher rate of speciation in the Indo-Pacific. In this group the species generally are of medium size and are moderately robust; they are usually marked with axial or zigzag color lines, or are of a distinct solid color. Apertural white lines, so common in certain species of Littorinas.s. and particularly of Austr- olittorina, Nodilittorina and Granulilittorina are usually absent or diffuse in Littoraria. All species probably produce a pelagic egg capsule, although observations on reproductive habits have been made on only a few. Possible evidence of a common lineage is given by the presence in most species of a basal flap on the penis; the flap contains no penial glands or “klebdrusén” as it does in other littorines. Synonymy— 1834 Littoraria Griffith, and Pidgeon, The Animal Kingdom, vol. 12, Mollusca and Radiata, Index, p. 598, pl. 1, fig 3. Type-species by monotypy: Littoraria pulchra ‘Gray’ (Sowerby, 1832) [= Littorina zebra (Donovan, 1825)]. Key To Principle Recent Species of Indo-Pacific Littoraria Leo ohell various ly Color patterned. sca. au io ad oo waves vd Soe ew hw hE a eS Ee Ld 2 Shell without color pattern, plain pinkish gray .............. coccinea p. 05-301 22 CC orumella wenerally: Dine to ViOlet -<..66 saad seGede evades Der ve Oe a4 woes ee 2M OMUeIIe WANTS ~ a soca ke ad ote ay Ace ew, bese Oe Od Owe OS ak: ace era hh gig d ews 4 3a. Color pattern of brown undulate fammules or spots 3b. Color pattern diffuse, whorls encircled with faint gray band 3c. Color pattern of rather straight, closely-spaced brown axial lines 4. Color pattern a “herring-bone”’ effect, shell, white apertural band present .. . re eee undulata p. 05-298 . mauritiana p. 05-304 .. kraussi p. 05-306 spire less than half the length of 586 ogee 3 eee ace Be praetermissa p. 05-307 4. Color pattern of closely-spaced brown punctations, spire more than half the length of shell, no white apertural band Ee ee ee pintado p. 05-297 [05-297] 436 Littoraria Littorina undulata Gray, 1839 (Pls. 325, 332, 333) Range—From Madagascar to western Polyne- sia; apparently rare in Hawaii; not reported from southeastern Polynesia nor from east Africa. Remarks—Due to its variability some forms of Littorina undulata are rather easily confused with several other Indo-Pacific species: L. scabra and L. coccinea, L. mauritiana and L. kraussi. From L. scabra it differs markedly in the anatomy of external reproductive structures, a large penile basal flap being present in undulata, whereas in scabra the entire base of the penis is thickened; scabra reproduces ovoviviparously while undul- ata produces a pelagic egg capsule; scabra usu- ally inhabits mangroves and other wooden struc- tures such as pilings, but undulata is usually Joseph Rosewater Littorinidae found on rocks; the shell of undulata is smaller (24 mm.) at maturity than scabra (over 40 mm.); although color patterns may appear superficially similar, upon close examination it may be noted that in scabra axial color markings are applied in a mosaic, the interruptions caused by the deeply incised spiral sculpture, while in undulata the markings appear continuous. The relationship is closer between undulata and the three other species, coccinea, kraussi and mauritiana. The features of all four are somewhat similar, and indeed they are considered members of the same subgenus, Littoraria. In the case of these species differences from undulata are more subtle and involve character of and presence or absence of color pattern, sculptural differences, relative proportions and geographical ranges (see species descriptions). x 12.6 mm.). Figs. 3,4. Lectotype of L. tenuis Philippi from Ticao Island, Philippines (BM(NH) 1968276; 14.7 x 8.4 mm.). Figs. 5,6,7,8. L. acuminata Gould from Mangsi Island [Philippines] (figs. 5,6: Holotype, USNM 5638; 8.9 x 4.8 mm.; figs. 7,8: paratype, USNM 612325, 6.8 x 3.7 mm.). Figs. 9-16. L. undulata from various localities: figs. 9,10. Pulau Stupai, Mentawai Ids., S.W. Sumatra (USNM 655218; 17.8 x 10.8 mm.). figs. 11,12. Pulau Siburu, N. of Sipora, Mentawai Ids., $.W. Sumatra (USNM 654710; 12.2 x 7.3 mm.); figs. 13,14. Bikini, Marshall Islands (USNM 585104; 20.05 x 11.8 mm.). figs. 15,16. Hien- ghene, New Caledonia (USNM 666080; 17.3 x 10.05 mm.). Fig. 17. Lectotype figure of L. columna ‘Jonas’ Philippi, from Abbildungen und Beschreibungen Conchylien, vol. 3, pl. 6, fig. 15 (about 18 x 11 mm.). [05-298] November 30, 1970 The western Atlantic Littorina nebulosa (Lamarck) is the analogue of the Pacific L. undu- lata. Habitat—Lives intertidally on rocky shores and commonly clusters in rock crevices during day- light hours, becoming active at night and crawl- ing about on the rocks. Often associated with L. coccinea (Gmelin) where their ranges overlap. Description—Shell reaching 23.9 mm. (about .9 inch) in length, turbinate in shape, average obesi- ty about .58 (24 specimens range from .50 to .64) moderately thick in structure, imperforate, and sculptured with spirally incised lines, overall microscopic wavy spiral threads and fine, but rather regular oblique axial lines of growth. Ex- ternal color very variable: from a mottled yellow- ish gray to banded dark-brown, most often with continuous to interrupted undulating brown col- or markings; spire whorls often with white flam- mules on upper half, solid brown below; aperture light yellowish brown to dark yellowish brown sometimes with outside color markings showing through; columella usually grayish violet but sometimes lacking this color and being white, edged with brown. Base not flattened, separated from upper part of body whorl by a weak keel at periphery. Whorls 5-9, usually rather well round- ed. Spire less than half the length of shell, convex, produced at an angle of about 58°. Aperture broadly oval; outer lip thin in young individuals to moderately thick in adults; inner lip strongly produced, hardly concave, oblique, glazed with a grayish violet callus, pinched where it joins outer lip at base of columella. Suture impressed. Sculp- ture consisting of from 7-10 spiral striae on spire whorls, persisting on entire surface of body whorl where as many as 16 may be present above weak keel, and about the same number below; on body whorl somewhat weaker secondary striae may occur between primary ones especially at suture and near periphery. Entire surface covered with microscopic, closely-spaced wavy spiral threads not easily detectable in worn specimens. Axial sculpture consisting of rather regularly-spaced, oblique growth lines which upon crossing spiral sculpture produce a weak reticulation. Opercu- lum corneus, paucispiral. Periostracum not evi- dent in specimens examined. Nuclear whorls partially decollate in all specimens examined; remaining portions smooth and glassy; first 2-3 post-nuclear whorls light-brown and sculptured with spiral striae. Radula typically littorinid (2- 1-1-1-2). INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 437 Animal darkly pigmented on upper surface of tentacles, snout and foot. Verge with a large flap at its base, long and slender, but constricted about half its length and with a delicate translucent distal extremity. Seminal duct deeply folded. Re- production oviparous; egg capsules observed to contain single ovum; capsule simple, convexo- convex, with flotation “‘skirt.”” Capsule diameter ranging from .23-.25 mm., ovum diameter .09-.11 mm. Development undoubtedly pelagic; well-de- veloped veligers remaining in capsules on fifth day (Eniwetok-February) after shedding of cap- sule; pelagic stage probably long lived. Measurements (mm)— length width no. locality whorls 93.9 11.9 8+ Pulau Nias, $.W. Sumatra 21.8 12.0 6+ Guadalcanal, Solomon Islands 19.8 10.6 6+ Guadalcanal, Solomon Islands 19.7 11.6 6+ S.W. tip Sanding Id., Mentawai Ids. 18.0 10.3 6+ Guadalcanal, Solomon Islands 17.4 TT 5+ Bikini Atoll, Marshall Islands L5.7 9.9 6+ Bikini Atoll, Marshall Islands 14.5 7.8 7+ Pulau Nias, $.W. Sumatra 11.3 6.5 8 Pulau Nias, S.W. Sumatra 10.9 6.1 8 Pulau Nias, S.W. Sumatra Synonymy— 1839 Littorina undulata Gray, The Zoology of Captain Bee- chey’s Voyage—in His Majesty’s Ship Blossom, Mol- lusks, p. 140 (no locality given; not figured); type locality here designated: Okinawa, Ryukyu Islands; Lectotype BM(NH) 1968362, British Museum (N.H.). Plate 333. Littorina (Littoraria) undulata (Gray) Fig. A. Radula (upper scale is 0.1 mm.). Fig. B. Penis (lower scale is 2 mm.). Fig. C. Egg capsule from above. Fig. D. Egg capsule from side to show flotation “skirt” (capsule diameter about 0.23 mm.); all from Eniwetok, Marshall Islands. 438 Littoraria 1846 Littorina tenuis Philippi, Proceedings of the Zoological Society of London (1845) p. 141 (Ticao Island, Phil- ippines); 1847 Abbildungen und Beschreibungen Conchylien, vol. 3, p. 18, pl 6, fig 8; lectotype and two paralectotypes, British Museum (N.H.) 1968276 1847 Litorina columna ‘Jonas’ Philippi, Abbildungen und Beschreibungen Conchylien, vol. 3, p. 14, Litorina pl. 6, fig. 15 (no locality given). 1849 Littorina acuminata Gould, Proceedings of the Boston Society of Natural History, vol. 3, p. 84 (Mangsi [Mangsee] Id. [Balabac Strait, Philippines]); 1852, U.S. Exploring Expedition, Vol. 12, p. 200, pl. 14, fig 239 a, b, Holotype: USNM 5638. 1878 Litorina scabra concolor Weinkauff, Systematisches Conchylien-Cabinet, vol. 2, part 9, p. 37, pl. 4, fig. 11 [indicated in text]; in part (figs 8-10 are L. scabra) (Indo-Pacific). 1885 Littorina undulata contracta Nevill, Hand List of Mol- lusea in the Indian Museum, Calcutta, part 2, p. 143 (Andamans). 1885 Littorina undulata sulcatula Nevill, ibid.; p. 144 (Ascen- sion Island, Pacific [Ponape, Caroline Ids.]); refers to Reeve, 1857, Conchologia Iconica, vol. 10, Littorina, pl. 13, fig. 67a [occasional specimens approach this degree of rugosity]. 1885 Littorina conica subintermedia Nevill, ibid., p. 150 (Port Canning and False Point [Bengal, India]). Types—From the 3 syntypes of L. undulata in the British Museum (NH), catalogue number BM(NH) 1968362, the one most closely approxi- mating Gray’s measurements and description is here designated as lectotype (length 22.8, width 12.6 mm.; see our pl. 332 figs. 1, 2). Gray listed no type locality, but there are few places where the Blossom touched land within the range of this species (see Beechey, 1831; Rosewater, 1968). One such place, Okinawa, Ryukyu Islands, is here designated as the type locality. The speci- men of L. tenuis Philippi BM(NH) 1968276, figured in Abbildungen und Beschreibungen z0° 40° 60° 80° \oo° _ 1z0° Joseph Rosewater Littorinidae Conchylien pl. 6, fig 8, is here designated lecto- type (14.7 x 8.4 mm.; see our pl. 332 figs. 3, 4). The type specimen of L. columna Philippi may be in the Berlin Museum. The types of L. con- tracta, subintermedia and sulcatula Nevill should be in the Indian Museum, Calcutta. The holotype of L. acuminata Gould is in the U.S. National Museum (USNM 5638). The location of the type of L. concolor Weinkauff is unknown to me. Records—MADAGASCAR: Ambodifototra, Ile Ste. Marie; Ile aux Nattes, S. of Ile Ste. Marie; Faty, 13 mi. N. of Tulear (all MCZ); MAURITIUS: Pt. d’Espy N.E. of Poste de Flacq; E. of Souillac on Savanne River (both ANSP); SEYCHEL- LES: Anse a la Mouche, Mahe Island (YPM); MALDIVES: South half Kendikolu Island, Miladummadulu Atoll; btwn. Mafilefuri and Maro Islands, Fadiffolu Atoll; Dunidu Island, N. of Male, North Male Atoll; Ari Atoll (all USNM, ANSP); Hitadu Id., Haddumati Atoll (YPM); Wala Island, South Nilandu Atoll; N.W. tip of Gan, Addu Atoll (both ANSP). INDIA: Vengurla, N. of Goa; Goa; Khumpta (Kumta), N. Kanara (all USNM); Cochin, Malabar (MCZ, AMS); Cape Comorin; Remen Point, west side Pamban Pass, btwn. Gulf of Mannar and Palk Strait (both ANSP, USNM): Mandapam Camp (USNM); Madras (ANSP, MCZ). CEYLON: shore, Ft. Frederick, Trincomalee (ANSP, YPM); Weligama Bay; Pointe de Galle; Galle (all ANSP); Columbo (USNM, MCZ, NMW, SMF); Delft Island; Kankesanturai; W. of Kankesanturai; Merissa Village (all ANSP). MALAYSIA: Pulau Anyut, Malac- ca Strait, just S.W. of town of Malacca (USNM). THAILAND: Goh Huyong, Similan Islands; Goh Phi Phi (both USNM). FORMOSA: (ANSP). JAPAN: Higo (Hiigo Shima) (USNM); Hachijo Island, 275 mi. S. of Tokyo, Honshu (ANSP); Kago- shima (USNM): Osima, Osumi (USNM; MCZ). RYUKYU ISLANDS: Okinoerabu-shima (ANSP); 1 mi. N. of Shana Wan, Okinawa Island (USNM); Nago, Okinawa Island (BPBM, MCZ); Awa (ANSP). PHILIPPINES: Many locali- ties, including the following islands: Batanes; Babuyanes; Luzon; Burias; Mindoro; Panay; Cebu; Bohol; Mindanao; Basilan; Palawan (all USNM). SULU ARCHIPELAGO: Jolo Island; Simaluc Island, Tataan Islands, Tawi Tawi Group; Marongas Island, Jolo (all USNM). BORNEO: Sipitang; West Marudu Bay; Taganak (all USNM). CELEBES: Macassar (MCZ). MOLUCCAS: Karakelong Island, Talaud Islands (MCZ); Morotai, Halmahera Group, (USNM); Kasiroeta Is- e' UNDULATA | COCOE - KEELING ATOLLS Ow 20°

% 4}: MARQUESAS 1S TUAM, | OTU Arey | Se emt i] ss geeeRlines au Ie ‘Ss. os ae UST Ra, ; |S. | NEW ZEALAND | Morio Theresa 1 n Qi 140° 160° ; 180° 60° 40° Plate 334. Geographical distribution of Littorina (Littoraria) undulata (Gray). [05-300] November 30, 1970 land; Salangadeke Island, N. Loloda, W. of Halmahera Island (both MCZ); Ceram (USNM); Hajara, Manipa Island (MCZ). SUMATRA: Pulau Melila, S. of Udjung Batu, Banjak Islands (USNM); Pulau We (MCZ, ZMA); Pulau Boenta, off Acheh Head; Pulau Nias (both USNM); Padang (MCZ); Pulau Sibu- ru, N. of Sipora; Veeckens Bay, South Pagi Island; Pulau Stupai, north edge Sanding Island, Mentawai Islands; south- west tip of Sanding Island, Mentawai Islands (all USNM); Marong (ANSP). JAVA: Pelabuhan Ratu, Praeger (MCZ; USNM); Welkomst Bay, Banten (USNM); Pemeungpeuk, nr. Garut, S.E. coast of Java (MCZ). BALI (AMS). FLORES ID. (ZMA). COCOS KEELING: North Keeling, near Pulo Selwa; Cocos Lagoon; “Coral Pits,’ Klapa Tuju, West Island (all USNM). WESTERN AUSTRALIA: Broome; Ridell’s Beach, 2 mi. S.E. of Gantheaume Point, Broome; Gantheaume Point, Broome (all ANSP); S. end South Pasco Island, Barrow Island Group, off Onslow (WAM, USNM). NEW SOUTH WALES: nr. Wollongong (USNM); Port Stephens (AMS). QUEENS- LAND: Heron Island, Capricorn Group (AMS); Cape Cleve- land, nr. Townsville (USNM); Palm Island (AMS); Brook Island (ANSP); Flying Fish Point; Russell River; Fitzroy Island; nr. Cairns; Lizard Island (all AMS); Green Island, nr. Cairns (MCZ); Cooktown; Murray Id., Torres Strait (both AMS). NEW GUINEA: Hollandia; Aitape; outlet of Kalveng River, Gusika, 13 mi. N. of Finschhafen (all MCZ); Huon Gulf (USNM); Collingwood Bay (AMS); Yule Island (USNM; BPBM:; ANSP; MCZ): Island 1/2 mi. S.S.W. OF Sowek, Soepiori, Schouten Islands (MCZ); Biak; N.E. end of Noekori Island; E. Noesi Isle, Mios Woendi Atoll, Padaido Islands; Rainbawi Point, Japen Island; S.E. entrance to Wooi Bay, Japen Island; Montoewoeri Point, Koeroedoe Island; Abroeki Island, Maransabodi Island, Aoeri Islands, Geelvink Bay (all ANSP). ADMIRALTY IDS.: Koruniat Island (ANSP). BIS- MARKS: New Ireland (MCZ, USNM); Matupi Island, Rabaul, New Britain (AMS). SOLOMONS: Woodlark Island (USNM); Harapa, Shortland Island (MCZ); nr. mouth of Lunga River, N. shore Guadalcanal (MCZ, ANSP); Point Purvis, Florida Island; Ugi Island (both USNM); Makira Harbor, San Cris- tobal (AMS). SANTA CRUZ IDS.: Tinakula (AMS). NEW HEBRIDES: Pakea, Banks Group (AMS). LOYALTIES: Li- fou (USNM; AMS); Maré (MCZ). NEW CALEDONIA: N. of Hienghene (AMS); Koe Reef, 2 mi. $.S.E. Touho (ANSP); Isle of Pines (MCZ); Ilot Maitre, about 4 mi. S.W. of Noumea; (ANSP): Ilot Amédie (AMS); La Roche, Bourail; Gatope Island, Voh (both ANSP). LORD HOWE ISLAND: Norfolk Island (both AMS). FIJI: Point N. of Namu, NNE Viti Levu (MCZ); Irvines, nr. Malaqereqere, S.W. Viti Levu (USNM); Lakemba (BPBM). MARIANAS: Saipan (ANSP; USNM;:NMW); Tinian (MCZ); Tumon Bay, Guam (BPBM). PALAUS: Koror, Malakal Dock; Eil Malk (both ANSP); An- gaur (BPBM, MCZ). HELEN REEF: N. end Helen Islands (ANSP). CAROLINE ISLANDS: Tomil Harbor Yap Island (USNM); Ulithi Atoll (ANSP); Elato Atoll (USNM); Dublon, Truk; Ponape (both BPBM); Kusaie (AMS, BPBM, MCZ, USNM); Kapingamarangi (USNM). MARSHALL ISLANDS: Eniwetok; Bikini; Kwajelein; Arno; Jaluit; Ebon Atoll (ALL USNM). GILBERTS: Abaiang (MCZ); Maiana; Tabiteuea (both BPBM); HAWAIIAN ISLANDS: Kailua, Oahu (BPBM); Ka- milo, Kau; Keaukaha, Hilo, both Hawaii (both USNM). LINE Islands; Palmyra (USNM). JOHNSTON ISLAND (USNM, BPBM). ELLICE ISLANDS: Nui; Vaitupu; Nukualailai (all USNM). WALLIS ISLANDS: Nukuhifala; btwn. Luanna and Fungalei Islands (both USNM). HOORN ISLANDS: Futuna (USNM). PHOENIX ISLANDS: Canton Island (USNM). SA- MOA: Salelesi Village, Upolu Island (ANSP); Pago Pago, Tutuila (BPBM; USNM). TONGA: Niuafoou Island; Niutoua, Tongatapu; Laulea Reef, nr. Fatuma, Tongatapu (all USNM); Velitoa; Tongatapu; Nomuka, Ha’apai (both BPBM). INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 439 Littorina coccinea (Gmelin, 1791) (Pls. 325, 335, 336) Range— Cocos-Keeling Islands to Eastern Polynesia, including Hawaii. Remarks—The name coccinea is derived from the Latin ““coccineus” meaning “red like a berry” and seems to be a misnomer in the case of adult specimens which are largely whitish. There is, however, always a blush of brownish orange coloration which shows through and the aperture is intensely colored. Young individuals show more of the color which perhaps prompted Gme- lin to adopt Martyn’s appellation and, in addition, Martyn’s plate shows an exaggeratedly red- colored invididual. Although evidencing no out- standing characteristics the species is easily rec- ognizable and in the center of its range, Microne- sia, probably is the most common littorinid pres- ent (see Demond, 1957). Habitat— Lives intertidally on rocky shores and commonly clusters in rock crevices during daylight hours, becoming active at night and crawling about on the rocks. Often associated with L. undulata Gray where their ranges over- lap. Description—Shell reaching 25.8 mm. (slightly over 1 inch) in length, elongate- to conic- Plate 335. Littorina (Littoraria) coccinea (Gmelin). Figs. 1,2. Specimen from Aitutaki, Cook Islands, showing diffuse color-banding (USNM 684817; 14.3 « 8.1 mm.). Figs. 3,6. Lectotype figure, from T. Martyn, Universal Con- chologist, vol. 2, pl. 68 [external figs]. Figs. 4,5. Specimen from Vitilevu, Fiji, nearly white with little evidence of external color or banding (USNM 531795; 22.7 x 14.0 mm.). [05-301] 440 Littoraria turbinate in shape, average obesity about .60, 21 specimens ranging from .56-.65, females slightly more obese than males), older individuals rela- tively thick in structure, imperforate, and sculp- tured with spirally incised lines, overall micro- scopic, wavy, spiral threads and irregular, ob- lique lines of growth. External color white, lack- ing patterned color markings except for occasion- al circular white blotches on early whorls; with brownish orange to strong brown within the aperture and showing through on lower portions of early whorls, on center of body whorl and on the base of shell. Base only slightly flattened, separated from upper part of body whorl by a weak to moderate keel at the periphery. Whorls 6-9, moderately rounded. Spire less than half the length of shell, convex, produced at an angle of about 60°. Aperture roundly oval; outer lip thin in young individuals to moderately thick in adults; inner lip concave, curves obliquely posteriorly, glazed with the brownish orange callus, pinched where it joins outer lip at base of columella. Suture distinctly impressed. Sculpture consisting of from 7-10 spiral striae over entire surface of early spire whorls, but often becoming indistinct on penultimate and body whorls where the first 1-5 striae only may be visible; first subsutural stria the strongest, pro- ducing a shouldered effect. Entire surface of shell covered with microscopic, closely-spaced wavy spiral threads not easily detectable in worn speci- 0.4mm t =) Plate 336. Littorina (Littoraria) coccinea (Gmelin), from Eniwetok, Marshall Islands. Fig. A. Radula (upper scale = 0.1 mm.) Fig. B. Penis (lower scale is 5 mm.) Fig. C. Two views of egg capsule (diameter of capsule about 0.24 mm.; diameter of egg about 0.09 mm.). Joseph Rosewater Littorinidae mens. Axial sculpture consisting of occasionally coarse but usually fine oblique lines of growth. Operculum corneus, paucispiral. Periostracum not evident in specimens examined. Nuclear whorls partially decollate in all specimens ex- amined; remaining portions smooth and glassy; first 2-3 post-nuclear whorls brownish orange and sculptured with spiral striae. Radula typically littorinid (2-1-1-1-2). Animal darkly pigmented on upper surfaces of tentacles, snout and foot. Verge long and slender, with a swollen enlargement at its base. Reproduc- tion oviparous; pelagic egg capsules observed to contain single ovum; capsule simple, convexo- convex, with flotation “skirt.” Capsule diameter 0.24; ovum diameter .09 mm. Development un- doubtedly pelagic; veliger stage reached in 2-3 days (Eniwetok-February) after shedding of cap- sules; some larvae swimming free.at that time. Measurements (mm)— length width no. whorls locality 25.8 15.0 6+ Keaukaha, Hilo, Hawaii 23.0 14.3 5+ Bikini Atoll, Marshall Ids. 20.4 12.0 9 Futuna, Hoorn Ids. 18.1 10.5 8+ Makatea, Tuamotu Ids. 15.3 9.2 8+ Makatea, Tuamotu Ids. 14.3 9.3 9 Makatea, Tuamotu Ids. 12.7 74 7 Huahine, Society Ids. 11.2 6.3 8+ Moorea, Society Ids. 9.8 5.8 7 Moorea, Society Ids. 7.2 4.2 6+ Moorea, Society Ids. Synonymy— [1788 Limax coccinea Martyn, Universal Conchologist, Vol. 2, plate 68; rejected work, non-binomial. ] 1791 Helix coccinea Gmelin, Systema Naturae, edition 13, vol. 1, part 6, page 3651 (New Zealand [Tahiti, Society Islands, here corrected]); refers to Martyn, Univ. Conch. 2, t. 68, f. ext [restricted to external figures; the figure on left (apertural view) here designated as lectotype]. 1832 Littorina obesa Sowerby, Genera of Recent and Fossil Shells, vol. 32, no. 37, fig, 6 [fig 6 here designated as lectotype] (South Sea Islands) 1839 Littorina limax Gray, the Zoology of Captain Beechey’s Voyage in His Majesty’s Ship Blossom p. 139 (Society Islands); refers to Martyn, Univ. Conch. [vol. 2, pl. 68, left external figure here designated as lectotype]. Types—Gmelin based the name Helix coccinea on the figures in Martyn’s Universal Conchologist (vol. 2, pl. 68, external figures). The location of Martyn’s collection is not known and therefore we must resort to his figures to distinguish this fortunately distinctive species. The left hand fig- ure, apertural view, is here designated as lecto- type of Gmelin’s Helix coccinea. The type locali- ty, given by Gmelin as New Zealand is here corrected to Tahiti, Society Islands, a likely place [05-302] November 30,-1970 for Martyn’s figured specimen to have been col- lected. The location of type specimens of L. obesa Sowerby and of L. limax Gray are also unknown; they were not found in the collection of the British Museum (N.H.) during a recent visit there (1968). For L. obesa, Sowerby’s figure 6 is here designated as lectotype; for L. limax, Martyn’s left external ne pl. 68 of the Universal Con- chologist, vol. 2, is here designated as lectotype. Records—FORMOSA (ANSP). RYUKYU ISLANDS: Kikai- ga-shima; Okinawa (both USNM). PHILIPPINES: Sabtan Island; Santo Domingo, Batan Island, both Batanes Group; Cujangan Island; Candaraman Island, Balabac (all USNM). INDONESIA: Karakelong Island, Moluccas (MCZ). CO- COS-KEELING ISLANDS: North Keeling (USNM). AUS- TRALIA: Murray Island, Torres Strait; Lizard Island, (both AMS); Green Island, nr. Cairns (MCZ); Herald Cays, off Cairns; West Cay, Diamond Islets; Russell Island; Flying Fish Point, all Queensland (all AMS), NEW GUINEA: 5 mi. N.W. of Rani Island, Biak Island, Schouten Islands, (ANSP); Hollandia (MCZ). BISMARCKS: Malie Island, Lihir Islands, New Ireland (AMS). SOLOMONS: Stirling Isle, Treasury Islands (USNM); Simbo Island (AMS, ANSP); Ugi Island (USNM); Mortlock Ids. (Del. Mus. Nat. Hist.). SANTA CRUZ ISLANDS: Vanikoro (AMS). NEW HEBRIDES: Espiritu Santo Island (MCZ); Vate Island (Efate Island) (AMS); Tana Island (USNM). LOYALTIES: Uvea Island (USNM); Lifou Island (AMS). NEW CALEDONIA: N. of Touho (ANSP, USNM); Ilot Maitre $.W. of Noumea (ANSP). LORD HOWE ISLAND (AMS): FIJI: Cevu Island, Nandronga; Irvines, Nr. Malaqeregere, both Viti Levu; Fulanga, Lau Group (all USNM). MARIANAS: Saipan Island (ANSP, MCZ); Tinian (USNM, MCZ); Tumon Bay, Guam (BPBM); Apra Bay, Guam (USNM). PALAUS: S.E. Eil Malk; S.E. Auluptagel Island, Malakal Harbor, Koror (both ANSP). CAROLINES: Yap (BPBM); Mog Mog Ulithi (BPBM, eee: Ponape (MCZ); Kusaie, “Coral Islet’ Lele (BPBM); Kapingamarangi (USNM, INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 441 BPBM). MARSHALL ISLANDS (many localities on the fol- lowing atolls): Eniwetok; Bikini; Rongelap; Rongerik; Ujae; Pokak; Bikar; Taka; Uterik; Wotho; Likiep; Ailuk; Arno; ie Kwajalein; Ailinglapalap; Jaluit Atoll (all USNM). GIL- BERTS: Abaiang Island (MCZ); Abemama Island (USNM): Tabiteuea (BPBM). HAWAIIAN ISLANDS: Kaaawa, USNM); Koko Head, western side, both Oahu (BPBM): Hilo, (MCZ, USNM); Kapoho, (BPBM); Pohoiki, all Hawaii USNM). LINE ISLANDS: Kingman Reef (BPBM); Palmyra Island (ANSP, USNM, AMS, MCZ, BPBM); Washington Island BPBM); Fanning Island (BPBM, AMS); Flint Island (USNM, ANSP); JOHNSTON ISLAND (USNM, MCZ, BPBM). EL- LICE ISLANDS: Nanumea; Nui (both USNM); Funafuti AMS, MCZ, USNM); Nukulailai (USNM); Rotuma (BMNH). WALLIS ISLAND: Nukuhifala (USNM). HOORN _IS- LANDS: Futuna (USNM). HOWLAND ISLAND (BPBM). BAKER ISLAND (BPBM). PHOENIX ISLANDS: Canton Island (USNM). SAMOA: Apia (USNM, AMS); Saluafata reefs, both Upolu (ANSP); Pago Pago Harbor, (BPBM, USNM); Aunuu Island, both Tutuila (MCZ); Ta’u Island (BPBM, MCZ); Fakaofo Island (BPBM), Tokelau Ids. TON- GA: Niuafo’ou (USNM; BPBM); Lifuka, Ha’apai Group; Velitoa, (both BPBM); Popua Reef, Tongatapu (USNM). COOK ISLANDS: North tip Akitua, Aitutaki (USNM); Taun- ganui, Mauke (BPBM); Rarotonga (AMS); Avatiu Harbor to Motu Tou, Rarotonga; Mangaia (both USNM, MCZ). AUS- TRAL ISLANDS: Raeivavae, (USNM). SOCIETY ISLANDS: Fanui Bay; S. of Farepiti Point; Vaitape V illage all Bora Bora; around point from “Hipu”’; Bay Vaiore both Tahaa; Uturoa; Il6t Tipaemanu S.E. of Utoroa; Tevaitoa, all Raiatea; Point Teffaao, around reef, Huahine; Moto Fareone; Opunohu Bay (all USNM); Pareau Point, District of Teavaro (ANSP), all Moorea; Tiarei; btwn. Hamuta River and Fautaua River, Pirae; Patutoa (all USNM); Atiue, Punaauia (ANSP); 43 kin. N. of Pariauta, Hitiaa, (USNM); btwn. mouths of Maruapo and Hapaa Rivers, District of Punaauia (ANSP); 1 km. S. of Faaone River, Faaone; Motu Fenuaina, Tautira (both USNM) all Tahiti. TUAMOTUS: Tikehau; Vahitahi; Reao; Tureia; Makemo (all USNM); Takaroa; Niau; Toao; Amanu (all ANSP): Makatea; Fakarava; Raroia (all ANSP; USNM). GAM- BIER ISLANDS: Mangareva (MCZ; USNM: AMS). Oeno Island; Pitcairn Island (both USNM). HENDERSON IS- LAND (Smith, 1913). 60° 80° too® 120° 1408 160° 180° 160 - 140 120 100 90 | | oo ni | \ ee / a { 3 oh \t ~ POC CINEA| a i - COCCINEA Sy ANG peat ING ( YS - { -- — — & os _A 20 os ATOLLS NEW ZEALAND a | 1 a an Tcoe*d Ce | | GALAPAGOS | | | 1s | 1 MARQUESAS IS. it : Too* Bo" 180° Plate 337. Geographical distribution of Littorina (Littoraria)coccinea (Gmelin), [05-303] 442 Littoraria Joseph Rosewater Littorinidae Littorina mauritiana (Lamarck, 1822) (Pls. 325, 338, 339) Range— South Africa and Southwestern Indian Ocean Islands. Remarks—Littorina mauritiana is restricted in its distribution to the southwestern Indian Ocean. It has been confused quite often with similar appearing species from South Africa, Australia and New Zealand but is unquestionably distinct when its peculiarly smooth gray banded shell and diffuse markings are recognized. The species most commonly confused with L. mauritiana is L. unifasciata and its subspecies in Australia, New Zealand and Juan Fernandez, but it is easily distinguished on the basis of shell and animal characters. This species is closely related to Litto- rina coccinea of the Western Pacific and also L. kraussi of the Indian Ocean. Habitat—Intertidal on rocks. Description—Shell reaching 24 mm. (about 0.9 inch) in length, turbinate in shape; average obesi- ty about .59 (13 specimens range from .55 to .63), older individuals relatively thick in structure, imperforate; sculptured spirally with overall mi- croscopic wavy threads and axially with occa- sional lines of growth. External color pattern rather constant, consisting of a spiral band of gray on middle of body whorl and on anterior third of spire whorls; area anterior and posterior to band with indistinct wavy axial color lines; anterior color lines ending at another rather indistinct gray spiral band on the base of body whorl with only occasional subsequent, sometimes obliquely spiral, color markings; overall background color a yellowish gray. Aperture usually dark brown, often with outside color pattern showing through; with an internal basal white spiral band and occasionally a small one posteriorly inside outer lip. Whorls 6-8 but spire often eroded. Spire less than half the length of shell, convex, produced at an angle of about 57° (but varying to as low as 48°). Aperture oval; outer lip thin in young speci- mens to moderately thick in adults; inner lip concave, curving obliquely posteriorly, glazed with a brown to violet callus, pinched where it joins outer lip at base of columella. Suture im- pressed. Sculpture consisting of microscopic, closely spaced, wavy spiral threads not easily detectable in worn specimens. Axial sculpture consisting of irregular lines of growth which may Plate 338. Littorina (Littoraria) mauritiana (Lamarck) Figs. 1,2. Lectotype of P. mauritiana from “‘Tle-de’France,” (MHNG 1096/91-2, 22.8 x 13.2 mm.). Figs. 3,4. Paralectotype (MHNG 1096/91-5, 12.5 x 7.4 mm.). Figs. 5,6. Lectotype of Littorina laevis Philippi, (BM(NH) 1968221, 18.4 x 11.4 mm.). Figs. 7,8. Holotype of Tricolia vitrea Deshayes from Bour- bon [Reunion] (MHNP, 6.1 x 4.2 mm.). Figs. 9,10. L. mauritiana from Pointe Fayette, E. Mauriti- us, (USNM 637330 [ex ANSP 273712] 19.8 x 11.1 mm.). Figs. 11,12. L. mauritiana from E. of Souillac, Mauritius, (USNM 637331 [ex. ANSP 274078] 15.8 x 9.4 mm.). (Figs. 1,2,7,8. R. Robertson photos; other photos by the author) have associated dark pigmentation. Operculum corneus, paucispiral. Periostracum not evident in specimens examined. Nuclear whorls partially decollate in all specimens examined; remaining portions smooth and unsculptured. Radula typi- cally littorinid. Animal darkly pigmented on upper surfaces of tentacles, snout and foot. Verge moderately long and slender, with a terminally swollen branch near its base, similar to L. kraussi. Reproduction unknown; [probably oviparous and involving pe- lagic capsules]. [05-304 ] November 30, 1970 Measurements (mm.)— length width no. whorls locality 24.0 13.3 7+ Mauritius PA P22 6+ Mauritius 19.9 EL 8+ Mauritius 19.6 12.4 5+ Madagascar 18.9 10.8 5+ Mauritius 18.1 11.4 7+ Reunion 17.5 il 6+ Mauritius 17.4 9.8 6+ Mauritius 14.1 8.7 5+ Mauritius 11.4 7.0 6+ Mauritius Synonymy— 1822 Phasianella mauritiana Lamarck, Histoire Naturelle des Animaux sans Vertebres, vol. 7, p. 54 (Ile-de-France [Mauritius]); Lectotype in Museum d Histoire Na- 1838 Phasianella mauriciana ‘Lamarck’ Potiez and Michaud, Galerie des Mollusques, Muséum de Douai, vol. 1, p. 311 (error for P. mauritiana Lamarck, 1822). 1846 Littorina laevis Philippi, Proceedings of the Zoological Society of London (1845), part 13, p. 140 (locality unknown [Mauritius here selected]); Lectotype in British Museum (N.H.); BM(NH) 1968221, 18.4 x 11.4 mm. 1847, Abbildungen und Beschreibungen Con- chylien, vol. 3, p. 10, Littorina, pl. 6, fig 6. 1847 Litorina mauritiana gracilior Philippi, Abbildungen und Beschreibungen Conchylien, vol. 2, p. 165, Littorina, pl. 3, fig 17b [right hand fig.] Probably is eastern Pacific species best known as L. modesta paytensis Philippi, 1847. 1863 Phasianella vitrea Deshayes, Catalogue des Mollusques de L’ Ile de la Réunion, (Bourbon) p. 76, pl. 8, fig 8 (Bourbon; Holotype in Muséum d Histoire Naturelle, Paris, 6.1 x 4.2 mm.). Types—There are 5 syntypes of Phasianella mauritiana in the Lamarck Collection of the Plate 339. Littorina (Littoraria) mauritiana (Lamarck) from Pointe Fayette, E. Mauritius (ANSP 273712) Fig. A. Radula (upper scale is 0.1 mm.) Fig. B. Penis (lower scale is 5 mm.) INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 443 Muséum d Histoire Naturelle, Geneva, represent- ing three different species of Indo-Pacific Lit- torinidae: L. mauritiana, L. kraussi, and L. uni- fasciata. The last cannot have occurred at Mauri- tius, which Lamarck clearly stated as type locali- ty, and L. kraussi does not fulfill both require- ments of Lamarck’s description: shell “bluish white” and columella “violet blue.” Syntypes 2 and 5 (MHNG 1096/91-2, 5) do fulfill these re- quirements and are here designated as Lectotype and paralectotype respectively of Phasianella mauritiana Lamarck (see our pl. 338, figs. 1-4) The specimen of L. laevis which Philippi fig- ured in Abbildungen und Beschreibungen Con- chylien vol. 3, pl. 6, fig 6, is in the British Museum (NH) BM(NH) 1968221, and is here selected as lectotype of that species (see our pl. 338, figs. 5,6). The holotype of P. vitrea Deshayes is in the Muséum d'Histoire Naturelle, Paris (see our pl. 338, figs. 7, 8). The type specimen of L. gracilior Philippi may still be extant in the Berlin Museum. Records—SOUTH AFRICA: Port Edward, Natal (ANSP). MADAGASCAR: Flacourt, Fort Dauphin (MCZ). SEYCHEL- LES: Praslin Island (Melville, 1909). MAURITIUS: S. side Tombeau Bay, W. Mauritius; E. Tamarin Bay, W. Mauritius; Pointe Fayette, E. Mauritius; E. of Souillac on Savanne River (all ANSP); Port Louis (MCZ). REUNION: Sainte Pierre (AMS). CHAGOS ARCHIPELAGO (Melville, 1909). Plate 340. Geographical distribution of Littorina (Littoraria) mauritiana (Lamarck) in the southwestern Indian Ocean. [05-305] 444 Littoraria Littorina kraussi new name Rosewater, 1970 (Pls. 325, 341) Range—East coast of Africa, the Indian Ocean islands and south India to Cocos-Keeling Atolls. Remarks—The familiar name Littorina glabra- ta Philippi, 1846, is unfortunately a secondary homonym of Paludina glabrata Pfeiffer, 1828 (=Littorina neritoides (Linné)), the Mediter- ranean species and, therefore, must be renamed. The species is renamed for F. Krauss from whom Philippi “stole” several South African species by publishing his manuscript names first. Littorina kraussi is closely related to L. mauritiana, undu- lata and coccinea. Habitat—On rocks near high tide line. Description—Shell reaching 21.8 mm. (about 0.9 inch) in length, turbinate in shape; average obesity about .58 (15 specimens range from .55 to .62); older individuals relatively thick in struc- ture, imperforate, and sculptured with spirally incised lines, overall microscopic wavy spiral Plate 341. Littorina (Littoraria) kraussi Rosewater. Figs. 1,2. Lectotype, L. glabrata [= L. kraussi] from Natal, South Africa (BM(NH) 1968220, 18.3 x 10.8 mm.). Figs. 3,4. Paralectotype, same BM number, 18.4 x 11.1 mm. Figs. 5,6. Tiladummati Atoll, Maldive Islands (USNM 672385, 17.7 x 10.5 mm.). Figs. 7,8. The same, a smaller specimen showing more intense coloration (11.4 x 7.4 mm.). Figs. 9,10. Zanzibar (USNM 89411, 14.7 x 8.9 mm.). Figs. 11,12. Near Poste de Flacq, Mauritius (USNM 637326, 15.5 x 8.7 mm.). Joseph Rosewater Littorinidae threads and fine rather regular oblique axial lines of growth. External color pattern rather constant, consisting of moderately regularly spaced, ob- lique, zigzagging, or chevron-shaped brown markings superimposed on a lighter background; (pale to grayish-yellowish pink ISCC-NBS num- bers 31-32) spire whorls above penultimate whorl often dark anteriorly and light posteriorly, par- ticularly noticeable in small individuals; aperture light to darker brown, sometimes with exterior color markings showing through; columella vari- able in color; ranging from dark brown through violet to nearly white with a brownish tinge. Base not flattened, separated from upper part of body whorl by a weak to moderate keel at periphery. Whorls 6-7 or 8 but spire often eroded. Spire less than half the length of shell, convex, produced at an angle of about 55°. Aperture oval; outer lip thin in young specimens to moderately thick in adults; inner lip concave, curving obliquely posteriorly, glazed with a brown, violet or nearly white callus, pinched where it joins outer lip at base of colu- mella. Suture impressed. Sculpture consisting of from 7-10 spiral striae on spire whorls and per- sisting onto body whorl where 10-11 may be present above the keel and about the same num- ber below; striae evanescent in some specimens, worn away or weak at center of whorls. Entire Zo* 40° : 60° 890° i YE Se FS ANDAME 1s NICOB/ 18. EYLON cocos | ee Reunion 80° Plate 342. Geographical distribution of Littorina (Littoraria) kraussi Rosewater in the Indian Ocean. [05-306 ] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 445 surface covered with microscopic, closely spaced, wavy spiral threads not easily detectable in worn specimens. Axial sculpture consisting of rather regularly spaced lines of growth. Operculum cor- neus, paucispiral. Periostracum not evident in specimens examined. Nuclear whorls partially decollate in all specimens examined; remaining portions smooth and glassy; first 1-2 post nuclear whorls light brown and usually sculptured with spiral striae. Radula typically littorinid and simi- lar to L. mauritiana. Animal darkly pigmented on upper surfaces of tentacles, snout and foot. Verge long and slender with a terminally swollen branch near its base, appearing very similar to L. mauritiana. Re- production unknown; [probably oviparous and involving pelagic capsules. ] Measurements (mm.)— length width no. whorls locality 21.8 11.9 6+ Mauritius 21.6 12.4 7+ Mauritius 19.5 11.6 T+ Mauritius Meo 10.5 7+ Zanzibar 16.9 9.3 6+ Madagascar 14.4 8.6 7+ Mombasa, Kenya 13.1 Uso 6+ Madagascar 12.5 6.9 7+ Mombasa, Kenya 10.2 6.3 7+ Mombasa, Kenya 9.3 D0 7 Madagascar Synonymy— 1846 Littorina glabrata Philippi, Proceedings of the Zoologi- cal Society of London (1845) p. 140 (Payta Peru; and Cape Natal [South Africa] here restricted to second locality). 1848, Abbildungen und Beschreibungen Conchylien, Vol. 3, p. 62. Litorina pl. 7, fig 5. Lectotype in BM (NH) 1968220; [Non Paludina gla- brata Pfeiffer, 1828=Littorina neritoides (Linnaeus, 1758)]. 1848, Krauss, Die Sudafrikanischen Mollus- ken, p. 103. Types—Although it was necessary to supply a new name for the secondary homonym, Littorina glabrata Philippi, 1846, the type specimens are the same for both L. glabrata and L. kraussi. A lectotype is here designated from among the four syntypes of L. glabrata in the British Museum (NH) (BM (NH) 1968220, length 18.3, width 10.8 mm.; see our pl. 341, figs. 1, 2). Records—SOUTH AFRICA: Miller's Point, False Bay (ZMA); Coffee Bay, 1 mi. S. Umtata River; East London, at mouth Nahoon River; Second Beach, Port St. John; Port Edward; Margate, 11 miles south of Port Shepstone; Isipingo (all ANSP; NMW): Durban; mouth of Umhlali River (both MCZ). MOZAMBIQUE: Ilha da Inhaca, Delagoa Bay (ANSP). TANZANIA: Kendwa Island, 4 miles E.S.E. of Dar-es-Salaam (MCZ); Bungi, Kiwani Bay; Changa; E. side Puopo, Tumbat Island (all ANSP). KENYA: Mombasa Island (ANSP; NMW). ADEN: (RNHL; BM(NH); MHNP). MADA- GASCAR: Anakao, 20.5 miles S. of Tulear; Soalary, 16 miles S. of Tulear; 1 mile N.W. of St. Augustin, 14 miles S.E. of Tulear; bay S. side Nosy Iranja, 32 miles S.W. of Nossi Bay; Nosy Tany Kely, 4 miles S. of Nosy Be; Pte. Fievre, S. Nosy Be; S. of Anivorano, 12 miles N. of Ambodifototra, W. coast Ile Ste. Marie; Ambodifototre, Ile Ste. Marie; S.W. shore Ie aux Nattes S. of Ile Ste. Marie; Flacourt, Fort Dauphin (all R.W. Foster; ANSP; MCZ). SEYCHELLES: St. Pierre Islet; Mahé Island (both YPM); Praslin Id. (NMW). MAURITIUS: S.E. side Tamarin Bay; S. side Tombeau Bay; !/4 mile S.W. of Cape Malheureux; Pte. Lafayette; 1 mile N.E. of Poste de Flacq; Pte. Vacoas, 3 miles S. of Mahebourg; near Savanne River E. of Souillac (all ANSP). REUNION: (MCZ).MALDIVE ISLANDS: Faro Islet reef, N.W. of Filadu Island, Tiladum- mati Atoll; between Mafilefuri and Maro Islands, Fadiffolu Atoll; Dunidu Island, North Male Atoll; Wala Island, South Nilandu Atoll (all ANSP); Hitadu, Had- dummati Atoll (YPM); Gan, Addu Atoll (YPM; ANSP); Ari Atoll (ANSP). CHAGOS ARCHIPELAGO: Egmont Atoll (Melville, 1909). INDIA: Goa (USNM). CEYLON: Point de Galle (ANSP; RNHL). COCOS-KEELING ISLANDS: Cocos lagoon, North Keeling (both USNM). Littorina praetermissa May, 1909 (Pl. 325, 343, 344) Range—Tasmania, Victoria and South Aus- tralia. Remarks—The “neglected” littorine is a dis- tinct and easily recognizable species inhabiting the southeastern Australian region. Considering the narrow range it occupies it is not surprising that it escaped attention until named by May. The Plate 343. Littorina (Littoraria) praetermissa (May) Figs. 1,2. Low Head, Tasmania; showing light-brown nu- clear whorls and distinctive color pattern (USNM 671215: 11.9 x 7.7 mm.). Fig. 3. Lectotype, original illustration from Papers and Proceedings Royal Society of Tasmania for 1908, pl. 6, fig. 3; Tasmania. Figs. 4,5. Port Fairy, Victoria; large specimen showing considerable wear and/or corrosion (USNM 637353: 16 x 11 mm.). [05-307] 446 Littoraria relationships of L. praetermissa appear to lie more closely with the species of the tropical Indo-Pacific than they do with the other Aus- tralian species, such as L. unifasciata, although the radula of praetermissa is similar in the nar- rowness of the central tooth. It may be dist- inguished at once from L. unifasciata by its coarser spiral sculpture, color markings, less deeply impressed umbilical crescent, and con- siderably more rounded whorls. The animals also differ in details of verge anatomy. Habitat—F ound in clusters on rock surfaces at and above the high water mark. (Macpherson and Gabriel, 1962). Description—Shell reaching 16.7 mm. (about .7 inch) in length, globose-turbinate in shape, aver- age obesity about .66 (23 specimens range from .61 to .71); moderately thick in structure, im- perforate, developing a rather narrow crescent- shaped area adjacent to columellar callus; sculp- tured with shallow and relatively widely spaced spiral striae and somewhat irregular axial lines of growth; shell surface rather uneven and bumpy. External ground color of shell grayish white, with usually prominent brown zigzag lines overall. Aperture medium brown with a white band at junction of outer lip and columella; with white markings around inner margin of outer lip. Base hardly flattened, separated from upper part of body whorl by a distinct to obsolete keel at periphery. Whorls 4-5, rounded. Spire consider- ably less that half the length of shell, convex, produced at an angle of about 72°. Aperture oval in outline; outer lip moderately thick, having its origin high on body whorl above keel so that in some specimens keel enters aperture as a simu- lated parietal tooth; inner lip nearly straight. Columella yellowish to tannish white, moder- ately well excavated, with a low tooth-like swel- ling near junction with outer lip; also usually with a narrow crescent-shaped area on base ad- jacent to columellar callus. Suture distinct. Sculp- ture consisting of from 15-17 sometimes indis- tinct spiral striae on body whorl and from 7-11 on spire whorls; striae fairly distinct in young speci- mens to indistinct in old invidivuals; shell sur- face between striae (i.e. spiral cords) relatively flat; peripheral keel on body whorl moderately formed to obsolete. Shell surface overall rather uneven and bumpy. Axial sculpture consisting of irregular oblique lines of growth. Operculum corneous, rather thin, paucispiral, nucleus well delineated. Periostracum not evident in speci- mens examined. Nuclear whorls at least partially Joseph Rosewater Littorinidae decollate in all specimens examined; remaining portions light brown in color, smooth, about 3 in number. Radula littorinid (2-1-1-1-2) central tooth moderately wide; lateral large and heavy, outer marginal with 11 denticles. Available rather poorly preserved animals dark- ly pigmented on surface of tentacles snout and foot. Verge only moderately short and thick; hav- ing a simple, truncate basal flap. Nothing is known concerning reproduction and develop- ment of this species; probably oviparous, spawn- ing pelagic capsule. Measurements (mm.)— length width no. whorls locality 16.7 11.9 4+ South Australia [pA 9.5 5+ South Australia 14.7 9.5 4+ South Australia 14,1 9.6 4 Port Fairy, Victoria 13.8 8.5 4+ Port Fairy, Victoria 13.1 8.7 5 Low Head, Tasmania 12.0 7.8 5 Low Head, Tasmania a7 7.6 4+ Queenscliff, Victoria 9.6 6.3 4+ Port Phillip Heads, Tasmania 9.0 6.3 4 Queenscliff, Victoria Synonymy— 1909 Littorina praetermissa May, Papers and Proceedings of the Royal Society of Tasmania for 1908, p. 57, pl. 6, fig 3 (Tasmania; type deposited in Tasmanian Muse- um, Hobart: 15 x 11 mm; probable paratypes in Australian Museum, Sydney and in Museum of Com- parative Zoology). Records—SOUTH AUSTRALIA: Robe (NMW); Boat- swain’s Point, Nr. Robe (USNM, ANSP, MCZ); Encounter Bay; Adelaide (both ANSP). VICTORIA: Shelly Bay, San Remo (BM(NH)); Phillip Island (MCZ); Sandringham, Port Phillip (ANSP); Port Phillip (AMS); Queenscliff (USNM); Port Fairy (ANSP, USNM, AMS, NMW). TASMANIA: east coast, Tasmania (AMS, paratype); Burnie; Wineglass Bay; Cape Sorell; Opossum Bay; Sandy Cape (all AMS); Low Head (USNM, MCZ); Eaglehawk Neck (AMS, MCZ). Plate 344. Littorina (Littoraria) praetermissa (May) Fig. A. Radula (upper scale is 0.1 mm.). Fig. B. Penis (lower scale is 10 mm.); both from Port Fairy, Victoria, Australia, USNM 637353. [05-308 ] November 30, 1970 s oy NEw be HEBRIDES .- a , Ol: hs See Filis | | | | | | NEW ZEALAND fi é | | 00° 120° 140° 160° 160° Plate 345. Geographical distribution of Littorina (Littoraria) praetermissa (May) in southern Australia. Littorina pintado pintado (Wood, 1828) (Pls. 325, 346, 347) Range—Southwestern Indian Ocean, from South Africa to Mauritius; in the Western Pacific from Japan through the North Pacific Islands to Hawaii. Remarks—Littorina pintado evidences a very peculiar discontinuous distribution. It occurs ina rather isolated outpost in the southwestern Indian Ocean apparently skips the mainland of southeast Asia and the East Indies and becomes established again in the Western Pacific from whence it is distributed more uniformly eastward to Hawaii. In the eastern part of its range it is recorded only from the North Pacific, but in the Indian Ocean it occurs far south of the equator. This sort of interrupted distribution is difficult to explain un- less the concept is accepted of a once more evenly distributed species that for unknown reasons has disappeared from the center of its former range. The possibility of long range rafting or even ship transport cannot definitely be ruled out however. The species is apparently quite hardy and there is a note with two specimens in the USNM collec- tion indicating that they were brought from Ha- waii to Los Angeles, California, where they lived for a year without seawater; an almost identical observation was made by Edmondson (1946). A darker subspecies, L. pintado schmitti occurs at Clipperton Island. An apparent analogue of L. pintado is L. pullata Carpenter of the tropical eastern Pacific. Littorina tessellata Philippi of the INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 447 tropical western Atlantic, L. cingulifera Dunker, and L. cincta Quoy and Gaimard of New Zealand initially appear similar to pintado but upon closer examination are quite distinct. Habitat—Lives intertidally on rocky shores and commonly clusters in rock crevices during day- light hours, becoming active at night and crawl- ing about on rocks. Observed to be associated Plate 346. Figs. 1-12. Littorina (Littoraria) pintado pintado (Wood); Figs. 13-16. L. (Littoraria) pintado schmitti (Bartsch and Rehder). Fig. 1. Lectotype of Turbo pintado Wood, from Sandwich Isl.; a rather attenuate specimen (BM(NH) 1968368, 17 mm. length.) Fig. 2. Lectotype of Littorina serialis Eydoux and Souleyet, from iles Sandwich (MHNP, 18.6 x 11.8 mm.). Figs. 3,4. Lectotype of Litorina ambigua Philippi from Insulae Sandwich (BM(NH) 1968314, 15.9 x 9.7 mm.). Figs. 5,6. Specimen from Hilo, Hawaii (USNM 339401, 15.5 x 9.7 mm.). Figs. 7,8. Specimen from Bikini Atoll, Marshall Islands (USNM 585105, 13.4 x 7.6 mm.). Figs. 9,10. Specimen from Ani Jima, Bonin Islands (USNM 621892, 15.2 x 9.1 mm.). Figs. 11,12. Specimen from East of Souillac, Mauritius (USNM 637354, 12.2 x 7.3 mm.). Figs. 13,14. Holotype of Littorina schmitti Bartsch and Rehder, from Clipperton Island, eastern Pacific; note dark coloration in this male specimen (USNM 472547, 12.2 x 7.6 mm.; this is specimen figured in original description). Figs. 15,16. Paratype of L. schmitti Bartsch and Rehder, from Clipperton Island, eastern Pacific, female specimen, somewhat corroded, but showing dark coloration (USNM 472546, 14.7 x 8.8 mm.). [05-309] 448 Littoraria Joseph Rosewater with L. coccinea and L. undulata at Eniwetok and Bikini Atolls, Marshall Islands (personal ob- servations; USNM records) and with L. picta in Hawaii (Whipple, 1965; Struhsaker, 1966). Description—Shell reaching 20.6 mm. (0.8 inch) in length, rather conic-turbinate in shape, average obesity about .60 (18 specimens range from .57-.63); older individuals moderately thick in structure, imperforate, and sculptured with spirally incised lines, overall microscopic wavy spiral threads and oblique axial lines of growth. External color yellowish to purplish gray with an overall pattern of closely spaced reddish-brown streaks or spots; the latter usually arranged in spiral series and only seldom appear axially coor- dinated; the center of body whorl and anterior half of spire whorls often darker in color. Aper- ture dark brown, the interstices of outer color pattern showing through as light lines inside outer lip. Base hardly flattened, separated from upper part of body whorl by a rather weak keel at the periphery. Whorls 5-9, moderately rounded. Spire less than half the length of shell, convex, produced at an angle of about 62°. Aperture oval; outer lip thin in young individuals to moderately thick in adults; inner lip weakly concave to nearly straight, its inner margin a white crescent rimmed with dark brown, relatively stout posteriorly but pinched where it joins outer lip at base of colu- mella. Suture moderately impressed; whorls often slightly constricted below suture. Sculpture con- sisting of about 10-11 spiral striae over surface of spire whorls; from 10-14 striae on body whorl above apertural line and 16-20 below on the base; extra striae often inserted between main striae or Plate 347. Littorina (Littoraria) pintado pintado (Wood) Fig. A. Radula (upper scale is 0.1 mm.). Fig. B. Penis; note lack of branches, flaps or glands (lower scale is 2 mm.). Littorinidae in worn specimens sculpture may be partially obscured. Character of shell surface between striae variable, usually flat but may be raised rounded cords. Entire surface of shell covered with somewhat diffuse microscopic, closely spaced wavy spiral threads not easily detectable in worn specimens. Axial sculpture consisting of rather irregular growth lines which are often associated with dark axial pigmentation streaks. Operculum corneus, paucispiral. Periostracum not evident in specimen examined. Nuclear whorls partially decollate in all specimens ex- amined, remaining portions smooth and glassy; first 2-3 post-nuclear whorls dark brown and sculptured with spiral striae. Radula typically littorinid (2-1-1-1-2). Animal darkly pigmented on upper surface of tentacles, snout and foot. Verge simple, lacking lateral protuberances, grayish white. Reproduc- tion oviparous; pelagic egg capsule plano-convex in outline, measuring between .16 and .2 mm. in diameter, usually containing a single ovum mea- suring between .08 and .1 mm. (see Ostergaard, 1950; Whipple, 1965; for details of spawning and development see Struhsaker, 1966). Measurements (mm.)— length width no. whorls locality 20.6 1233 5+ Midway Id. 19.3 11.0 5+ Midway Id. 19.1 11.0 7+ Chichi Jima, Bonin Ids. 17.0 10.2 5+ Osima Osumi 15.9 10.0 7+ Hawaii 14.7 9.2 5+ Osima Osumi 14.2 8.8 6+ Mauritius 13.0 CD 7+ Hawaii 12.3 74 6+ Mauritius 10.1 6.2 6+ Osima Osumi Synonymy— 1828 Turbo pintado Wood, Supplement to the Index Testaceo- logicus, p. 20, pl. 6, Turbo fig. 34 (Sandwich Isl.); lectotype BM(NH) 1968368. 1839 Littorina tenebrata ‘Nuttall’ Jay, A Catalogue of the Shells in the Collection of John C. Jay, 3rd ed; p. 73 (Sandwich Isles) [nomen nudum]; 1847 [as Litorina tenebrata ‘Nuttall’] Philippi, Abbildungen und Be- schreibungen Conchylien, vol. 2, p. 203 [in synonymy of L. pintado Wood]. 1848 Litorina ambigua ‘Nuttall’ Philippi, Abbildungen und Beschreibungen Conchylien vol. 3, p. 62, Litorina, pl. 7, fig 6 (Insulae Sandwich); Lectotype in B.M. (NH) 1968314. 1852 Littorina serialis Eydoux and Souleyet, Voyage Autour du Monde sur La Corvette La Bonite, Zoologie, vol. 2. pl. 31, figs 34-36, (Iles Sandwich); lectotype in Museum d’Histoire Naturelle, Paris. 1882 Litorina pindata Philippi in Weinkauff, Systematischen Conchylien-Cabinet, vol. 2, part 9, pp. 63, 70 [error for L. pintado]. 1882 Littorina ambiqua Reeve in Weinkauff, ibid., p. 70 [error for L. ambigua]. [05-310] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 449 ° ° ° ° nN e T 1 ——+ 7 } 3 | oO | 1 cay \ ae Be ' — 2 eee) = y 6 ' io ' i 1 L \ £ ! 8 1 wy ' ' f | 1 £ 4 4 — _ a | = 4 ' if \ 4 ees q Lay “ ae 5 a & ® z ' o 1 ae ee “ S ! ! —>— 4 3 ' z - } ¢_ Fe 1 | o ’ H 8 aan f° de s rae || | tl i 1 SF 8. 18 8 i RP «gieg tS N \ : : - 2 §| oy, an | . G18 ‘ oo eae & 2 x ° Eel He ieee lo Q et et.) lo Co — - —— 18 2 tar Xa ea cy . of } OI ge@2 . ba | $2 a4 Lt a ant z H or. oe 20) 1 Jor ge Le | 5 ee roy . H zZ ° 5 : 34 CO‘k 3 4 3 ® 7 | 2 wy. —j— — wd iD 2 4 e 6 Zum |2 4 rss VE n> 3 2’ eS 227 4 g SI z oll eto Ae as | aw ae Fea oe 3a i a ~ wo 2 =" | = 4) % | Ee apie en ‘e e SEYCHELLES 1S. 40° ° o ° “ a Plate 348. Geographical distribution of Littorina (Littoraria) pintado pintado (Wood) in the North Pacific Ocean and in the southwestern Indian Ocean and of the subspecies, L. pintado schmitti (Bartsch and Rehder), from Clipperton Island, east- ern Pacific. Types—From 17 syntypes of Turbo pintado Wood from the Gray collection, the largest is here designated lectotype (see our pl. 346 fig. 1; length 17 mm.; BM(NH) 1968368. The figured speci- mens of L. tenebrata ‘Nuttall’ Philippi and L. ambigua ‘Nuttall’ Philippi, are here designated lectotypes of these species. The location of the figured specimen of L. tenebrata is not known, but that of L. ambigua is believed to be in the British Museum (NH) (BM(NH) 1968314; 15.9 x 9.7 mm.; (see our pl. 346 figs. 3, 4). The lectotype of L. serialis Eydoux and Souleyet, here designat- ed, is in the Muséum d Histoire Naturelle, Paris (see pl. 346 fig. 2; 18.6 x 11.8 mm.). Records—SOUTH AFRICA: Margate, 11 mi. S. Port Shep- stone, Natal, (ANSP). MOZAMBIQUE: Punta Abril, Delagoa Bay (ANSP). MADAGASCAR: S. of Anivorano, 12 mi, N. of Ambodifototra, W. coast He Ste. Marie; $.W.shore of Ile des Nattes, S. of Ile Ste. Marie; Flacourt, Fort Dauphin (all MCZ). MAURITIUS: Pointe Fayette; E.S.E. of Souillac (both ANSP); Souillac S. coast, E. of Souillac (ANSP; USNM). REUNION: (MCZ, NMW). JAPAN: Osumi Islands (USNM). RYUKYU ISLANDS: Shuri; Odomari, both Okinawa (both USNM). PHILIPPINES: Santo Domingo, Batan Island, Ba- tanes Group (USNM). BONIN ISLANDS: (ANSP, MCZ, NMW): Ani Jima; Chichi Jima (both USNM). MARIANAS: Agrigan; W. Pagan (both USNM); Saipan (ANSP, MCZ, USNM); Tinian (MCZ; BPBM; USNM). PALAUS: Angaur (ANSP; MCZ; USNM; SMF). WAKE ISLAND: (USNM; BPBM: ANSP). MARSHALL ISLANDS: Eniwetok; Bikini; Rongelap Atolls (all USNM). HAWAIIAN ISLANDS: Mid- way (USNM, MCZ, ANSP); Namokulaa, Eastern Islet, Mid- way Atoll; Pearl and Hermes Reef, Grass Island; Lisianski Island (all BPBM); Laysan (USNM, BPBM); Gardner Island; La Perouse Pinnacle, French Frigate Shoals (both BPBM); Necker Island (BPBM; USNM); Nihoa Island (BPBM); Kama- lino Bay, Niihau Island (USNM); and many localities on Kauai, Oahu, Molokai, Maui, Lanai, Kahoolawe and Hawaii (see map; ANSP; BPBM; USNM, MCZ). JOHNSTON IS- LAND: Sand Island (USNM, MCZ, BPBM). Littorina pintado schmitti (Bartsch and Rehder, 1939) (Pl. 346 figs. 13-16) Range—Known only from Clipperton Island. Remarks—Hertlein) and Emerson (1953) pointed out that a number of marine mollusks which are distributed mainly in the Indo-Pacific have been found on Clipperton Island, a locality which also has strong eastern Pacific faunal affin- ities. Littorina schmitti was described by Bartsch and Rehder as being related to L. pintado. The degree of relatedness is considered here to be definitely of subspecific rank, the very distinct differences between pintado and schmitti appa- rently having been caused by geographic isola- tion. Habitat—Living intertidally on rocks. [05-311] 450 Littoraria Joseph Rosewater Littorinidae Description—Largest available specimen 18.5 mm. (about 0.7 inch) in length, average obesity about .58 (20 specimens range from .54 to .64), somewhat shorter than L. pintado, similarly shaped and sculptured, but with darker colora- tion, the individual color spots tending to be larger and to coalesce, so that many specimens appear to have a dark background with light spots instead of the reverse as in L. pintado. Spiral striae usually 7-10 on spire whorls and on body whorl above apertural line, indistinct on the base; shell often marked with white spiral lines, both externally and within aperture; white lines often coincide with striae. Inner lip nearly straight when viewed from aperture; when viewed from aspect of outer lip, columella is thickened and pronouncedly convex at its midpoint. Measurements (mm.)— length width no. whorls locality 18.5 10.1 5 Paratype 18.4 9.9 4+ Paratype lrg 10.5 4+ Paratype 15.7 9.0 5 Paratype 14.9 9.0 6 Paratype 13.9 8.2 5 Paratype 13.0 7.8 5 Paratype 12.6 7.4 7 Paratype 12.2 7.6 6+ Holotype 10.3 6.6 5 Paratype Synonymy— 1939 Littorina schmitti Bartsch and Rehder, Smithsonian Miscellaneous Collections, vol 98, no. 10 (Publica- tion 3535), p. 9, pl. 2, fig 4 (Clipperton Island); Holotype: USNM 472547; original measurements of holotype (11.8 x 8.2 mm.) different from those ob- tained here: 12.2 x 7.6 mm. Records—Clipperton Island: on rocks, south of landing place (Holotype); east side, reef flat (both USNM),. Littorina sundaica Altena, 1945 (Pl. 349, figs. 1, 2) Range—Java, Indonesia. Remarks—The “Sunda” littorine is an ap- parently quite distinct species with a rather nar- row geographic distribution according to avail- able records, having been reported only from western Java. Due to its small size, however, it may be overlooked easily and when more collec- tions are made in nearby areas of Indonesia, the species may be found more widespread. The dark-brown color, few spirally incised lines of sculpture and single white spiral color band visi- ble internally and externally, together with its rather narrow shell outline are about the only distinctive characteristics of this species, yet they serve to distinguish it from any other known littorine. The shell of L. sundaica bears a slight similarity to the species assigned to Rissolittorina Ponder, 1966, R. alta Powell, and mariae Ten- ison-Woods, especially due to the spiral white band on the base of the shell of all three. How- ever, Rissolittorina appears to be a cool water group whereas L. sundaica obviously is a tropical species. Habitat—Not definitely known, but probably on shore rocks. Description—Shell reaching 8.5 mm. (about 5/16 inch) in length, elongate-narrow-turbinate in shape, average obesity about .55 (4 specimens range from .53 to .57); shell moderately thick in structure for its size, imperforate, smooth, except for a few spirally incised lines on body whorl and fine growth lines. Overall color of shell nearest to dark reddish brown (ISCC-NBS color number 44) becoming somewhat lighter brown in a diffuse band near suture and aperture edge; with a nar- row, white spiral color band on base below per- iphery also showing in aperture. Base hardly flattened, body whorl subcarinate at periphery. Whorls 5-7, very slightly rounded. Spire more than half length of shell, convex, produced at an angle of about 42°. Aperture subquadrate; outer lip moderately thick, inner lip nearly straight to only slightly concave, glazed with a brown callus, slightly pinched at junction with outer lip. Suture distinct; whorls may be slightly shouldered. Sculpture consists of 2-3 weak incised spiral striae just above periphery and 1-3 below it on base; fine axial and spiral growth lines are visible microscopically. Periostracum not evident. Oper- culum corneus, paucispiral. Nuclear whorls de- collate in all specimens examined. Radula, animal and reproduction unknown. Measurements (mm.)— length width no. whorls locality 8.5 4.5 7+ (holotype) Tjilaoet Eureun, Java 2 3.8 5+ “Java” 6.0 3.4 5+ Kaledjetan, Bantam, Java 5.5 3.0 6+ “Java” Synonymy— 1945 Littorina (Melaraphe) sundaica Altena, Zoologische Mededeelingen, Leiden, vol. 25, p. 151, fig 2, (Tji- laoet Eureun, south coast of Java); holotype in Rijks- museum van Natuurlijke Historie, Leiden, no. 52038. Records—JAVA: (MCZ, USNM); Kaledjetan, Bantam (USNM); Tjilaoet Eureun, (Holotype, RNHL). [05-312] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 451 Littorina acutispira E. A. Smith, 1892 (PI. 349, fig. 3; pl. 351, fig. 1) Range—Known only from coasts of southern Queensland and New South Wales, Australia. Remarks—This and the preceding species, L. sundaica Altena, are placed provisionally in the a aa ads Plate 349. Figs. 1,2. Littorina (Melarhaphe) sundaica Altena. Holotype, from Tjilaoet Eureun, S. coast of Java (Leiden Museum no. 52038, 8.4 x 4.4 mm.). Fig. 3. Littorina acutispira E. A. Smith. Lectotype from Port Jackson, New South Wales, Australia (BM(NH) 91.11.6.216-225, about 7 x 4 mm.). Figs. 4,5. Littorina lucida Yokoyama. Unique holotype, from Pliocene of Japan (Geological Institute, Imperial University of Tokyo, 5 x 3 mm.). Figs. 6,7. Littorina kozaiensis Nomura and Onisi. Unique holotype, from Lower Miocene of Japan (Saito H6-on Kai Museum, Sendai, Japan, Reg. no. 21762, 11 x 8 mm.). Figs, 8,9. Littorina adonis Yokoyama. Unique holotype, from Pliocene of Japan (Geological Institute, Imperial University of Tokyo, 6 x 4 mm.). subgenus Littoraria. More definite placement may be possible upon examination of fresh ana- tomical material, which has not been available for the present study. The general appearance of L. acutispira is littorinoid, but certain features, such as a tendency to form an umbilicus and the yellow bordered aperture which is dark in its interior, resemble some of the Fossaridae, notably F. atratus C. B. Adams of the tropical eastern Pacific. Because of the generalized nature of Lit- torina, it is not surprising that some species should resemble more generalized species of other fami- lies. The light-colored apertural edge is also seen in L. africana and there is a tendency toward this character in L. unifasciata. Habitat—‘Common in rock pools” (Smith, 1892). Description—Shell reaching 7 mm. (about 0.3 inch) in length, elongate-turbinate in shape, aver- age obesity about .57 (7 specimens range from .51 - .63); moderately thick in structure for its size, tending to become umbilicate, sculptured with weak spiral striae and irregular, coarse axial lines of growth. External color variable, from light yellowish brown to dark brown, usually with a wide spiral band of brown to gray encircling body whorl and anterior 2/3 of spire whorls; speckled on the body whorl with whitish spots. ° ° ° \oo0° 120 140 : 160 vi |) Mochy/o \ BONIN j 1S | . Iwo Jima - Moreus fe Sse eS = ese tibiae pce a eee eae ne “MARIANA | Mek is | / Soipon © ge PHILIPPINE * Guam ov or? Is. 2 Eniwetok ate, Yap, \ Uithi | i - . 7 = Wi - a re is fe Be eh ot 7 Ponape wey : CAROLINE “1s. ° Bee “Helen Rt : NEW per CALEDONIA _ “8 _ Norfolk ITISPIRA — dN — —EE 100° 120° 140° 160° Plate 350. Geographical distribution of the species, Littorina (Littoraria) sundaica Altena, in Indonesia, and of L. acutis- pira E. A. Smith, in eastern Australia. [05-313] 452 Littoraria Aperture dark reddish-brown within and on colu- mellar lip; outer lip with a broad white band and a white stripe revolving into it. Base slightly flattened; body whorl weakly carinate. Whorls 4-6, rounded. Spire more than half the length of shell, convex, produced at an angle of from 45- 50°. Aperture oval; outer lip moderately thick, inner lip slightly concave, glazed with a brown callus, flattened, and pinched near junction with outer lip; when umbilicus is present there is a trough just outside columellar callus. Suture dis- tinct, whorls hardly shouldered. Spiral sculpture consists of well-spaced, weak spiral striae over surface of whorls; axial sculpture of irregular axial lines of growth. Periostracum not evident. Operculum corneus, paucispiral. Nuclear whorls decollate in all specimens examined. Radula, ani- mal and reproduction unknown. Measurements (mm)— length width no. whorls locality 7 3.5 6 lectotype 5.6 3.4 5 All from Port Jackson, 5.6 3.1 5 New South Wales, 5.0 2.6 5 Australia 4.5 2.5 5+ 4.3 2.7 4+ 3.8 2.2, 4 Synonymy— 1892 Littorina acutispira E. A. Smith, Proceedings of the Zoological Society of London, part 4, (1891), p. 487, pl. 40, fig 3 (rock pools, Green Point, Watson Bay, Port Jackson, New South Wales); lectotype BM(NH) 91.11.6.216-225. Types—One of the 14 syntypes of L. acutispira Smith (BM(NH) 91.11.6.216-225) is here chosen as lectotype for the species (see our pl. 349 fig. of 7 X 3.0 mim,), Records—AUSTRALIA: Gladstone; Caloundra, both Queensland (both AMS); Port Macquarie; Catherine Hill Bay; Middle Harbor (all AMS); Port Jackson, all New South Wales (AMS; MCZ; USNM). Littorina infans E, A. Smith, 1892 (Pl. 351, fig. 2) Range—New South Wales and Queensland, Australia. Descriptive Remarks—Littorina infans is in- cluded provisionally here although its small size and the lack of any real proof of anatomical relationships causes me to consider it more a doubtful species of Littorinidae. Final generic and possibly even family assignment must await further investigation. Joseph Rosewater Littorinidae Plate 351. Fig. 1. Littorina acutispira E. A. Smith, from Port Jackson, New South Wales, Australia (from original figure in Proc. Zool. Soc., London, for 1891, pl. 40, fig. 3; about 7 x 3.5 mm.). Fig. 2. Littorina infans E. A. Smith, from Port Jackson, New South Wales, Australia (from original figure in Proc. Zool. Soc., London, for 1891, pl. 40, fig. 4; about 3.5 x 2.3 mm.). Synonymy— 1892 Littorina infans E. A. Smith. Proceedings of the Zoologi- cal Society of London (1891) p. 488, pl. 40, fig. 4 (Green Point, Watson Bay, Port Jackson [New South Wales, Australia] “found in rocky pools washed by ordinary high tides” (Brazier); syntypes BM(NH); 91.11.6.226-235; original measurements 3.5 x 2.3 mm. Records—AUSTRALIA: NEW SOUTH WALES: Green Point, Watson's Bay, Port Jackson (Syntypes: BM(NH); AMS); Sussex Haven; Little Coogee Bay; Shark Island. QUEENS- LAND: Burleigh Heads; Caloundra; Marouchydore (all AMS). [05-314] November 30, 1970 Littorina kozaiensis Nomura and Onisi, 1940 (P1. 349, figs. 6, 7) Range—Lower Miocene of Japan. Remarks—This species was described as re- sembling L. adonis Yokoyama (see below), but as having a larger number of spiral grooves. The unique holotype (pl. 349, figs. 6, 7, a copy of the original illustration) offers little basis for com- parison with Recent species. Synonymy— 1940 Littorina kozaiensis Nomura and Onisi, Japanese Jour- nal of Geology and Geography, vol. 17, nos. 3 and 4, p. 191, pl. 19, fig 6 a,b. (Yosuibori, Simizu, Kozai- mura, Japan); holotype: Saito Ho-on Kai Museum, Register No. 21762; 11 x 8 mm. Littorina adonis Yokoyama, 1927 (Pl. 349, figs. 8, 9) Range—Pliocene of Japan. Remarks—Placed_ provisionally here in the subgenus Littoraria, this species resembles L. undulata, although the strong spiral sculpture is also reminiscent of Littorinopsis, i.e. L. scabra, etc, Unfortunately the outer lip and a portion of the body whorl of the type (pl. 349, figs. 8, 9) are missing and it is difficult to be sure of their exact shape. INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 453 Synonymy— 1927 Littorina adonis Yokoyama, Journal of the Faculty of Science Imperial University of Tokyo, section 2, vol. 1, part 10, p. 451, pl. 51, fig. 8, (Upper Musashino, Koyasu southern Musashi, Japan); (unique holotype in collection of Geological Institute Imperial Univer- sity of Tokyo: 6 x 4 mm.). Littorina lucida Yokoyama, 1927 (Pl. 349, figs. 4, 5) Range—Pliocene of Japan. Remarks—This species is from the same de- posit as L. adonis but lacks the deeply incised spiral sculpture. The type of lucida, although of approximately the same size as adonis is more slender. This type (pl. 349, figs. 4, 5) is also broken, but the columella is complete and the relationship with members of the subgenus Lit- toraria is more easily confirmed than was the case with L. adonis. Synonymy— 1927 Littorina lucida Yokoyama, Journal of the Faculty of Science Imperial University of Tokyo, section 2, vol. 1, part 10, p. 451, pl. 51, fig. 9 (Upper Musashino, Koyasu southern Musashi, Japan; unique holotype in collection of Geological Institute Imperial University of Tokyo: 5 x 3 mm.). [05-317] 454 Littoraria Joseph Rosewater Littorinidae [ These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence. ] [05-318] November 30, 1970 Subgenus Littorinopsis Morch, 1876 Type: Littorina angulifera (Lamarck, 1822) The subgenus Littorinopsis is a small group of mostly tropical species, typified by L. angulifera, which are usually ovoviviparous and which show a habitat preference for mangroves or shore vege- tation. Individual populations of some species, such as L. scabra and its subspecies L. angulifera, exhibit a wide range of variation probably due to the geographical isolation imposed by their meth- od of reproduction which tends to restrict gene flow. Nevertheless, within their ranges, these species are widespread possibly because their habitat offers opportunities for rafting of adults. Shells of species of Littorinopsis show strong development of spiral sculpture which in some may form multiple carinae (L. carinifera, L. INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 455 scabra) and there appears to be a tendency for dark coloration and axial color banding and spot- ting to become strongly developed also. In males the penis is well-developed, with an open but folded sperm duct, a thickened base and a nar- rower distal filament. There are three Recent and two Tertiary fossil species of Littorinopsis included here. The ex- tremely variable L. scabra forms some local pop- ulations which have been considered as species. Most of these appear to be only minor genetical or ecotypical variants, not worthy of even sub- specific distinction (see scabra synonymy). Two species closely related to L. scabra (L. carinifera and L. melanostoma) appear to have developed sufficient genetic constancy to be considered as separate species. Synonymy 1876 Littorinopsis Morch, Malakozoologische Blitter, vol. 23, p. 135 (type-species by original designation Littorina subangulata Lamarck [lapsus for angulifera]). 1887 Lamellilitorina Tryon, Manual of Conchology, vol. 9, pp. 230, 253 (type-species by subsequent designa- tion, Wenz, 1939: Littorina (Lamellilitorina) albicans Metcalfe [= L. scabra Linné]}). Key to Recent Species of Indo-Pacific Littorinopsis ie olemells lig Diack. 4, sono aad es eee ee Se Sew e nwa eRe melanostoma hy Columetla lip mot Blak ~ ssid yok Gee a exceed odd Rb he AG embod we Sow Hie PRE A aes 2 2. Two broad white bands revolving within aperture, shell carinate and with narrow fed axial lipes.OrspOts: 454 4. ha ceo hor 64K SSS Owe e TER carinifera 2. Aperture with many fine lines and spotted with brown inside and out ....... scabra [05-329 ] 456 _Littorinopsis Joseph Rosewater Littorina scabra scabra (Linne, 1758) (Pls. 325, 352, 353) Range—Pan-Indo-Pacific, from South Africa to southeastern Polynesia. Remarks—The “rough” Littorina, which is one meaning of the Latin name _ scabra (another is “scabby,” which would also fit a brown-spotted shell) is an extremely ubiquitous species through- out the Indo-Pacific wherever there are man- groves or even wood pilings. Its comparatively large size would appear to make it easily visible were it not well camouflaged on mangrove trunks and branches. The relatively long synonymy is a partial indication of its variability. The species produces an almost endless variety of color and sculptural forms from the “typical” dark brown mosaicly banded scabra to the whitish rather smooth albicans described by Metcalfe. The mor- phological diversity is probably due to the fact that scabra is ovoviviparous and produces closely related, inbred populations within which varia- tions or mutations may develop and are perpetu- ated. The same phenomenon takes place in the Atlantic subspecies, angulifera whose biology is similar to scabra (Lenderking, 1954; Struhsaker, 1966). Plate 352. Littorina scabra (Linné). Showing a range of shell variations. Fig. 1. Helix scabra Linné (Lectotype figure, from Chem- nitz, vol. 11, pl. 210, fig. 2074, about 35 x 21 mm.). Fig. 2. Littorina arboricola Reeve (Lectotype, from Sin- gapore; BM(NH) 1968321, 31.9 x 25.4 mm.). Fig. 3. Littorina fortunei Reeve (Lectotype, from China; BM(NH) 1968309, 16.2 x 7.6 mm.). Fig. 4. Litorina flammea Philippi (Lectotype, from China; BM(NH) 1968310, 16.8 x 7.7 mm.). Fig. 5. Littorina albicans Metcalfe (Lectotype, from Sara- wak, Borneo; BM(NH) 1968355, 18.8 x 10.8 mm.). Fig. 6. Litorina scabra articulata Philippi (Lectotype, from Mindanao; BM(NH) 1968354, 31.5 x 16 mm.). Fig. 7. Littorina cingulata Philippi (Lectotype, from “north coast of Australia”; BM(NH) 1968352, 19.4 x 10.7 mm.). Fig. 8. Littorina intermedia Philippi (Lectotype, from Ne- gros Id., Philippines; BM(NH) 1968353, 14.2 x 7.8 mm.). Fig. 9. Littorina luteola Quoy and Gaimard (Lectotype, from Port Jackson, Sydney, Australia; MHNP, 16.3 x 9 mm.). Fig. 10. Littorina newcombi Reeve (Lectotype, from Ha- waii; BM(NH) 1968308, 25.3 x 15.1 mm.). Fig. 11. Littorina novaehiberniae Lesson (Lectotype, from Port Praslin, New Ireland; MHNG, 24 x 13.8 mm.). Fig. 12 Littorina pallescens Philippi (Lectotype, from Min- danao, Philippines; BM(NH) 1968277, 22.1 x 12.9 mm.). Fig. 13. Littorina philippiana Reeve (Lectotype, from “Philippines”; BM(NH) 1968307, 29.6 x 16 mm.). Figs. 14,15. Littorina scabra rhodea Biggs (Holotype, from Bundar Abbas, Persian Gulf; BM(NH) 1958.6.13.23, ca. 11.5 x 7.5 mm.). Littorinidae Variations in L. scabra generally take the form of yellow and orange color mutants. It may also exhibit normal closely-spaced spiral sculpture to more widely-spaced carinae. These variations oc- cur throughout the geographic range of the species more or less randomly. One variation which appears to be more geographically limited occurs on the north coast of Western Australia and was named L. sulculosa Philippi (see pl. 325, fig. 3). Specimens resembling this form were collected from ground rocks in a mangrove swamp on Barrow Island, Western Australia in 1966, and are considered to be ecophenotypes of L. scabra, having a habitat different from the latter. Since forms intermediate between scabra and sulculosa were found at the same locality, sulculosa does not appear to have validity as a specific entity. Differences between L. scabra scabra and its Atlantic subspecies L. scabra angulifera are dif- ficult to enumerate as they are mostly qualitative: angulifera tends to be lighter in color overall, ranging more toward yellow and orange than the darker brown scabra; its markings are more dif- fuse than in most of the latter; the aperture of angulifera is narrower, more subquadrate and the columella tends to have a violet hue, while in Fig. 16. Littorina sieboldii Philippi (Holotype, from Japan; BM(NH) 1968278, 29 x 15.4 mm.). Figs. 17,18. Littorina sulculosa Philippi (Lectotype, from “north coast of Australia”; BM(NH) 1968279, 17.7 x 9.9 mm.). Fig. 19. Specimen from Arue, Tahiti (USNM 668338, 15.2 x 9.7 mm.). Fig. 20. Specimen from Mbweni, Zanzibar (USNM 604470, 15 x 7.3 mm.). Fig. 21. Specimen from near Mombasa, Kenya (USNM 215230, 19.9 x 10.5 mm.). Figs. 22,23. Specimen from Proserpine, N. Queensland, Australia (USNM 679498, fig. 22: 23 x 13.2 mm.; fig. 23: 23.4 x 13.2 mm.). Figs. 24,25. Specimens from Po Bui Id., Sandakan, North Borneo (USNM 233252, fig. 24: 20.8 «x 11.5 mm., fig. 25: 22.6 x 11.5 mm.). Figs. 26,27. Specimen from Barrow Id., Western Australia (USNM 691687, 16.1 x 9.7 mm.). Figs. 28,29. Specimen from Sekudu Id., Strait of Johore, Malaysia (USNM 660732, 10 x 6.1 mm.). Fig. 30. Specimen from Broome, Western Australia (USNM 637333, 20.2 x 11.6 mm.). Fig. 31. Specimen from Santubong, Sarawak, Borneo (USNM 671209, 15.1 x 8 mm.). Fig. 32. Specimen from Guam, Marianas (USNM 426452, 39.6 x 22.7 mm.). Fig. 33. Specimen from Flat Top Id., near Mackay, Queens- land, Australia (USNM 622988, 25.4 x 14.2 mm.). Fig. 34. Specimen from Proserpine, Queensland, Australia (USNM 679503, 24 x 12.4 mm.). [05-330] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina Plate 352. Littorina scabra (Linné). Explanations on opposite page. [05-331] 458 Littorinopsis Joseph Rosewater Littorinidae scabra the aperture is more broadly rounded and the columella whitish or stained with brown; in angulifera there is often a series of brown spots just within the edge of the outer lip, while in scabra the entire interior of the aperture shows the external color, although this is rather variable in both species. Sculptural characters are essen- tially similar and isolated specimens are difficult to distinguish without comparison. I consider the range of L. scabra to be limited to the tropical Indo-Pacific, that of angulifera the tropical east and west Atlantic. Some introductions of L. angu- lifera into the east Pacific have taken place in the vicinity of Panama and the notable subspecies L. scabra abberans (Philippi, 1846), having a dark brown-rimmed aperture, occurs occasionally in that area. A possible explanation for the wide range of variation in color of L. scabra, in addition to the fact that populations are more reproductively isolated than many other marine snails by reason of being ovoviviparous, is that it is, in effect, largely a tree snail. Extreme color variation is recognized within and between populations of such pulmonate tree snail genera as Polymita and Liguus. Clench (1968) stated that for tree snails “coloration is apparently non-selective as there must be little ocular predation.” When snails leave the ground and ascend trees, they are im- mediately free of much of the danger from attacks by ground-living invertebrates and mammals which under ordinary conditions may select them for the familiar subdued coloration usually evi- denced by many exposed land, freshwater and marine snails. It may be theorized, therefore, that in L. scabra color variation is not under the control of selective forces usually exerted upon other species of Littorinidae and is, therefore, freely expressed in many of its populations. Habitat—Usually found on the trunks and branches of mangroves and on the ground; may be present on dock pilings and on sea walls; not commonly found on shore rocks except in areas from which mangroves are absent. Description—Shell reaching 43 mm. (about 1"/16 inches) in length [occasionally larger, but often smaller], high-turbinate in shape, average obesity about .57 (15 specimens range from .50 - .67); relatively thin in structure, usually im- perforate, and sculptured with spiral striae be- tween which on the body whorl are raised, split spiral cords; body whorl subcarinate to carinate at periphery. Color and color pattern variable, typi- cally a mosaic of brown blotches arranged axially or zigzag or in no apparent pattern; occasional populations with individuals partly or entirely lacking brown pattern and ranging in color from reddish orange through medium brown to yellow and white. Base often moderately flattened, sep- arated from upper part of body whorl by a moder- ately to well developed keel at periphery. Whorls 6-9, well rounded. Ratios of aperture and spire length to length of shell varying from equal to one exceeding the other. Spire convex, produced at an angle of from 43-57° depending on population. Aperture large, rather broadly rounded in outline; outer lip rather thin even in mature individuals, curving out widely; inner lip usually concave posteriorly (above) slightly convex anteriorly and almost forming a tooth just above where it joins outer lip near base of columella. Apertural por- tion of columella heavy, usually white or with violet to brown stains. Interior of aperture show- ing same mosaic color pattern as exterior. Suture distinctly impressed, whorls often showing a weak but differentiated subsutural cord standing out from preceding whorl. Sculpture consists of from 10-12 weak to relatively strong spiral cords with intervening shallow spiral striae; cords may or may not be split by secondary striae. Under magnification close-spaced, fine, wavy spiral threads are present over entire surface. Axial sculpture consists of fine lines of growth; occa- sionally growth lines are well-developed and some specimens may have spaced axial “ribs;” in some specimens every other spiral cord is sup- pressed and shell develops strongly carinate ap- pearance. Operculum large, thin, corneous, pau- cispiral. Periostracum not evident. Nuclear whorls partially decollate in all specimens ex- amined; remaining portions light-brown, spirally sculptured, and appear ornate. Radula littorinoid (2-1-1-1-2). Animal large, darkly pigmented on surfaces of tentacles, snout and foot; ctenidium well-developed in both males and females, serv- ing in latter as partitioned “brood pouch,” oc- cupying most of dorsal internal surface of mantle cavity. Penis with greatly thickened base, having lateral thickened appendage with bulbous ex- tremity; penis becoming considerably narrower at tip. Sperm groove open but folded. Reproduction ovoviviparous; eggs released into mantle cavity where held in ctenidial “brood pouch;” usually released in late veliger stage; free-swimming lar- val life probably very short. Egg size about 0.11 mm. [05-332] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 459 Plate 353. Littorina scabra (scabra (Linné). Fig. A. Radula of specimen from Sanga Sanga Id, Sulu Archipelago, Philippines, ANSP 230667; upper scale is 0.1 mm. Figs. B,C. Penis of specimen from Eniwetok, Marshall Ids.; fig. B. view from posterior showing sperm duct (detail indicates deep duct) fig. C. anterior view; specimen relaxed with Propylene Phenoxytol; scale is 5 mm. Measurements (mm.)— length width no. whorls locality 43.3 25.3 6 Luzon, Philippines 42.1 23.4 7 Oahu, Hawaii OOet 22.6 7 Guam 38.2 21.4 8 Barrow Id., W. Australia 35.3 19.9 8 Barrow Id., W. Australia 33.7 18.0 a Luzon, Philippines 30.7 18.5 7 Buka Id., Gulf of Tomini, Celebes 27.4 16.4 7 Mauritius 24.1 14.4 Jordan River, Guimaras Philippines 23.6 11.8 8+ Formosa 20.6 11.6 8 Koh Tau, Thailand 19.8 12.1 7+ Formosa 17.4 10.0 7 Vengurla, India 14.1 7.8 7+ Changanoue Strait, Mombasa, Kenya 11.0 6.0 "i Buena Vista, Guimaras Id., Philippines Synonymy— [1705 Buccinum foliorum Rumphius, d’Amboinische Rariteit- kamer, p. 98, pl. 29, species Y; prelinnean.] 1758 Helix scabra Linne, Systema Naturae, ed., 10, vol. 1, p. 770; type locality here designated: Amboina, Moluc- cas; Lectotype here designated: specimen figured by Chemnitz, vol. 11, pl. 210, fig. 2074. 1791 Buccinum lineatum Gmelin, Systema Naturae ed. 13, vol. 1, part 6, p. 3493; refers to Knorr, Vergn, 3, pl. 14, fig. 4, here designated lectotype (no locality given). 1831 Littorina novaehiberniae Lesson, Voyage Autour du Monde, Coquille, vol. 2, part 1, p. 348 (Port Praslin, Nouvelle-Irlande [Solomon Islands]. Lectotype here designated, specimen ex. Lesson in MHNG: 24 x 13.8 mm). 1832 Littorina luteola Quoy and Gaimard, Voyage de |’Astro- labe, vol. 2, p. 477, pl. 33, figs 4-7 (Port Jackson, [near] Sydney [Australia]); lectotype here designated one of 7 syntypes in MHNP: 16.3 x 9.0 mm. 1846 Littorina intermedia Philippi, Proceedings of the Zoo- logical Society of London, part 13 (1845), p. 141 (here restricted to Jimamailan, Negros Id., Philippines); lectotype here selected BM(NH) 1968353: 14.2 x 7.8 mm.; 1847; Abbildungen und Beschreibungen Con- chylien, vol. 2, p. 223, Litorina pl. 5 fig. 8, (lectotype). 1846 Littorina intermedia punctata Philippi, ibid., (no locality given in 1846); 1847, ibid; Litorina pl. 5, fig 11 (Tahiti; Elizabeth Island; Natal; Red Sea). 1846 Littorina intermedia articulata Philippi, ibid., (Swan Point [Western Australia]); 1847, ibid. 1846 Littorina intermedia strigata Philippi, ibid., (Ji- mamailan, Negros Id. Philippines); 1847, ibid., Litorina pl. 5, figs 8-10. (Mergui). 1846 Littorina pallescens Philippi, ibid., p. 142 (Province of Cagayan de Misamis, Mindanao [Philippines]); lec- totype BM (NH) 1968277: 22.1 x 12.9 mm.; 1847, ibid., vol. 3, p. 10 [corrected page] Litorina pl. 6, fig 4. 1846 Littorina sieboldii Philippi, ibid., (Japonia); holotype in BM(NH) 1968278: 29 x 15.4 mm.; 1847 ibid., p. 9, Litorina pl. 6, fig 3. 1846 Littorina cingulata Philippi, ibid., (ad oram borealem Novae Hollandiae); lectotype BM(NH) 1968352: 19.4 x 10.7 mm.; 1847, ibid., p. 11, Litorina pl. 6, fig 5. 1846 Littorina sulculosa Philippi, ibid.; (in Ora boreali Novae Hollandiae); lectotype BM (NH) 1968279: 17.7 x 9.9 mm.; 1847, ibid., p. 18, Litorina, pl. 6, fig 10. 1847 Litorina scabra flammulata Philippi, Abbildungen und Beschreibungen Conchylien, vol. 2, p. 222, Litorina (Panay; Singapore) 1847 Litorina scabra articulata “Menke” Philippi, ibid.; pl. 5, fig. 4, (Mindanao here restricted); lectotype BM(NH)1968354: 31.5 x 16 mm.) 1847 Litorina scabra punctata Philippi, ibid., pl. 5, fig 5. (Masbate; Singapore). 1847 Litorina scabra suturalis Philippi, ibid., pl. 5, fig 7 (Kanguruh-Inseln, dem Meersbusen Georgs des Vier- ten; Canton) 1847 Litorina scabra lutea Philippi, ibid., pl. 5, fig. 6 (Mas- bate; Philippines, Canton) 1847 Litorina scabra rubra Philippi, ibid., refers to Chemnitz [vol. 11, pl. 210] fig. 2075 (Mindoro, Philippines; Canton) 1847 Litorina scabra ventricosa Philippi, ibid., pl. 5, fig 8 (China; Mindanao; Tonga Tabu; Neu Irland; Neu Guinea); not L. ventricosa Philippi, ibid., vol. 3, p. 15. 1847 Litorina fammea Philippi, ibid., vol. 3, p. 16, Litorina, pl. 6, fig. 21 (China); lectotype BM (NH) 1968310: 16.8 x 7.7 mm. 1847 Litorina sinensis Philippi, ibid., vol. 3, p. 52, Litorina pl. 6, fig 23 (China). 1852 Littorina albicans Metcalfe, Proceedings of the Zoologi- cal Society of London, part 19 (1851) p. 73 (Sarawak, Borneo); lectotype BM(NH) 1968355: 18.8 x 10.8 mm.; 1857, Reeve, Conchologia Iconica, vol. 10, pl. 9, figs. 44a, b. 1857 Littorina philippiana Reeve, ibid., pl. 5, figs 22 a,b (Philippine Islands); lectotype BM(NH) 1968307, 29.6 x 16 mm. 1857 Littorina arboricola Reeve, ibid., pl. 6, fig 27 a,b (Sin- gapore); lectotype BM(NH) 1968321: 31.9 x 25.4 mm. 1857 Littorina newcombi Reeve, ibid., pl. 6, fig 28 a,b (Sand- wich Islands); lectotype BM (NH) 1968308: 25.3 x 15.1 mm. 1857 Littorina fortunei Reeve, ibid., pl. 9, figs 42 a,b China); lectotvype BM(NH) 1968309; 16.2 x 7.6 mm. [05-333] 460 Littorinopsis Joseph Rosewater Littorinidae 1871 Litorina strigata Lischke, Malakozoologische Blatter, vol. 18, p. 148 (Nagasaki [Japan]); type: Academy Science, Leningrad?; 1871 Japonische Meeres- Conchylien, vol. 2, p. 73, pl. 5, fig. 22. 1871 Melaraphe (Littorina) blandfordi Dunker, Malakozoo- logische Blatter, vol. 18, p. 150 (Rockhampton [Aus- tralia]); type in Berlin Museum? 1878 Litorina scabra concolor Weinkauff, Systematisches Conchylien-Cabinet, vol. 2, part 9, p. 37, pl. 4, figs 8-10 [not fig. 11 as indicated in text = L. undulata] (Indo-Pacific) [in part]. 1878 Litorina scabra minor Weinkauff, ibid., pl. 4, figs. 16-18 (Indo-Pacific); refers to L. intermedia Philippi and to L. newkombi (sic) Reeve. 1878 Littorina newkombi ‘Reeve’ in Weinkauff, ibid., pp. 37, 38 [error for L. newcombi Reeve]. 1885 Littorina scabra tenuis Nevill, Hand-List of Mollusca in the Indian Museum, part 2, p. 146 (Arakan [Burma]); type in Indian Museum, Calcutta?; not L. tenuis Philippi, 1846. [=L. undulata]. 1885 Littorina pallescens? erronea Nevill, ibid., p. 148 (Sin- gapore); type in Indian Museum, Calcutta? 1885 Littorina filosa subcingulata Nevill, ibid., p. 149 (Port Jackson [Australia]); type in Indian Museum, Calcut- taP 1885 Littorina conica delicatula Nevill, ibid., p. 149 (Port Canning and False Point (Bengal); type in Indian Museum, Calcutta?) 1885 Leptopoma (?) ardouinianum Heude, Memoires con- cernant I’Histoire naturelle de ]’Empire Chinois, Cahier 3, Notes sur les Mollusques terrestres de la vallee du Fleuve Bleu, p. 95, pl. 25. figs. 8, 8a (ad rupes Tonquinenses in portu dictu A-long) 1900 Littorina philippina von Martens, Biologia Centrali- Americana, Land and Freshwater Mollusca, p. 584; emendation for L. philippiana Reeve, 1857. 1958 Littorina (Melaraphe) scabra rhodea Biggs, Journal of Conchology, vol. 24, no. 8, p. 272 (Bundar Abbas [Persian Gulf]); holotype BM(NH) 1958.6.13.23, ca. 11.5 x 7.5 mm.) Types—In the original description of Helix scabra Linné refers to the Museum Ludovicae Ulricae, leading one to believe that a specimen under that name was present in that collection. Such a specimen is not contained, however, in the “MLU” of Odhner’s unpublished list [1953], nor is H. scabra represented in the Linnean Shell Collection in London. Furthermore, in the 12th Edition Linné did not refer to the Ulricae collec- tion under scabra. In order to clarify the concept of H. scabra, one of the figures referred to by Hanley (1855), is here designated as lectotype: Chemnitz, Conchylien Cabinet, vol. 11, pl. 210, fig. 2074 (see pl. 352, fig. 1). The specimen represented in that figure may be in the Zoologi- cal Museum, Copenhagen. The type locality for H. scabra is here designated as Amboina, Moluc- cas, as no locality was given by Linné. The location of holotypes and lectotypes of the other synonyms of scabra are mentioned in the synonymy where this information is known to me. Types of species described by Philippi in the “Abbildungen” are probably in the Berlin Muse- um except in those few cases where I designated as lectotypes the figured specimens found in the British Museum (NH); see Synonymy. The lectotype of L. albicans Metcalfe, men- tioned in the synonymy, which is the specimen figured by Reeve, is undoubtedly part of the original syntypic series, as Metcalfe’s collection was purchased by Reeve and parts of it were resold (The Athenaeum, No. 1906, May 7, 1864, p. 630). Records—SOUTH AFRICA: Port Alfred, nr. Grahamstown, Bathurst District, Cape Province (USNM, MCZ):; Durban; Isipingo, both Natal (both MCZ). MOZAMBIQUE: Inhaca Island, Delagoa Bay (NMW; ANSP). TANZANIA: Dar es Salaam; 1.5 mi. N.W. of Magogoni; Kendwa Island; Kun- duchi; Bagamoya; Tanga (all MCZ); Mboamagi, 9 mi. S. of Dar es Salaam (USNM); 2 mi. S. by W. of Chwaka, East Zanzibar; Bungi, Kiwani Bay, $.W. Zanzibar; 1 mi. N. of Chukwani, W. Zanzibar (all ANSP); Mbweni, 4 mi. S. of Zanzibar City; Bweju, S.E. Zanzibar (both USNM). KENYA: Mombasa Island (ANSP); Straits at Changanoue, 3.75 mi. from Mombasa, Kenya (USNM). MADAGASCAR: S. of Anivorano, 12 mi. N. of Ambodifototra, W. coast of Ile Ste. Marie; Ambodifotgtra, at causeway to Ilot Mandane, Ile Ste. Marie; S.W. shore Ie des Nattes, S. of Ile Ste. Marie; Soalary, 16 mi. S. of Tuléar; nr. Grottes de Sarodrano, 10 mi. S.W. of Tuléar; Tuléar (all MCZ); Nossi Be (ANSP; MCZ). SEY- CHELLES: Aldabra Island (Y.P.M.; USNM); Menai Island, Cosmoledo Atoll; Anse a la Mouche, Mahé Island; Northwest of Moyenne Island (all YPM). MAURITIUS: Point d’Espy, 1 mi. N. by E. of Poste de Flacq (USNM, ANSP); 3/4 mi. N. of Black River Bay (ANSP). MALDIVES: Male Harbor, Male Atoll; between Mafilefuri and Maro Islands, Fadiffolu Atoll (both ANSP); Gan, Addu Atoll (YPM; ANSP). INDIA: Bom- bay (USNM; MCZ): Vengurla, N. of Goa; Goa; Khumpta (Kumta), N. of Kanaru; Netravati River, Mangalore; Tuticorin; Adyar River estuary, Madras; Port Canning (all USNM); Cochin Harbor, Kerala; Rameswaren Island, Pamban, Palk Strait (both ANSP); Calcutta (MCZ). CEYLON: (MCZ, ANSP). BURMA: Victoria Point (MCZ). THAILAND: Ko Contee, Ranong (MCZ); Ko Sindarar Nua (Chance Island); Ko Phi Phi; Pulau Tanga, Butang Group (all USNM); Songkla (MCZ); Ban Tha Kham; Ko Phluai; Ko Tao (all USNM); Mutapone Island, Chumpon; Ban Hua Hin; Ko Si Chang, 40 mi. S. of Menam River; Bang Poo, Paknam; Ang Hin, Chol- buri Province; Bang Saen, Cholburi Province (all MCZ); Ko Chang; Ko Kut (both USNM). CAMBODIA: Kampot (MCZ). VIETNAM: Saigon (MCZ; MHNP). MALAYSIA: Penang (USNM, MCZ):; N.E. corner, Pulau Lumut, Port Swettenham; Pulau Besar, Malacca Strait, S.E. of town of Malacca; Cape Rachardo, Straits of Malacca; Sekudu Island, Strait of Johore (all USNM); Singapore (USNM, ANSP, MCZ). CHINA: Kiautschou, nr. Tsingtau, Yellow Sea; Spider Island, Fukien Province; Amoy (all USNM); Hong Kong (BM(NH), USNM, ANSP); Macao; Hainan (both ANSP. JAPAN: Awa (ANSP, MCZ). RYUKYU ISLANDS: (USNM). TAIWAN (USNM, ANSP, MCZ). PHILIPPINE ISLANDS (Many localities; see map). SUMATRA: Moro Id. (SMF); Pulau Melila, S. of Udjung Batu, Banyak Islands; S.E. coast of Pulau Nias; Pulau Bai, Batu Group (all USNM); Padang (MCZ); Pulau Siburu, N. of Sipora; S.W. tip of Sanding Island, Mentawi Islands (both USNM); Tapanuli Bay, Sibolga (ZMA). JAVA: Men- scheneter Island (MCZ, ANSP, USNM); Kaledjitan, Bantem (USNM); Djakarta (USNM, ZMA). BALI: Koeta Beach (USNM). BORNEO: Santubong, Sarawak (USNM, MCZ); Kudat Bay (ANSP, USNM); Tajong Aru, Jesselton (USNM); W. Marudu Bay (USNM, ANSP); Po Bui Island, Sandakan; Karamuntig, Sandakan; sandy plain, Sandakan Bay; Bohay- dulong Islands (all USNM). CELEBES: Dago Bay, Sangihe Islands; Manado (both MCZ); Likupang; Papajato River; Bukabuka Island, Gulf of Tomini; Limbe Island, Gulf of Tomini (all USNM); Wowoni Id., N. of Butung Island (MCZ); [05-334 ] November 30, 1970 Labuan Blanda Island, Butung Strait; Labuandata Bay, Gulf of Boni (Bone) (both USNM). MOLUCCAS: Talaud Islands; Morotai Island; Ternate; Kahatola Island, Loloda Islands, N.W. coast of Halmahera; Pajahi Bay, Halmahera Island; Kasirota Bay, Kasirota Island; Batjan Island; Oong Bay, Man- dioli Island; Buru Id.; Tengah Island, nr. Buru; Manipa Island (all MCZ); Amboina (ANSP); Kobroor, Aru Ids. (SMF); COCOS-KEELING ISLANDS: N.E. end Direction Island (USNM). AUSTRALIA: QUEENSLAND: Darnley Island (MCZ); Murray Island (AMS); Thursday Island, all Torres Strait; mouth of Norman River (both WAM); mud flats nr. Pilot Station, Karumba; Mapoon all Gulf of Carpentaria; Cape Sidmouth; Burkitt Island, nr. Port Stewart; Three Isles (all AMS): Cooktown (AMS, MCZ); Redlynch (MCZ); Cairns (USNM):; Green Island, off Cairns (USNM, MCZ, AMS); Atherton (MCZ); Flying Fish Point (AMS); Dunk Island; Palm Island (both ANSP, AMS); Cape Bowling Green (USNM): Bowen (AMS, MCZ); Hayman Island (AMS); Shute Harbor (USNM): Hamilton Island; Lindeman Island (both AMS); Proserpine (MCZ); Flat Top Island, nr. MacKay (USNM):; Percy Island, off Palmerston (MCZ); Yeppoon (AMS): Gladstone (USNM, ANSP); Port Curtis; Port Vernon; Pialba; Urangan (all AMS); Moreton Bay (MCZ); Sandgate, nr. Brisbane (ANSP, AMS); Stradbroke Island (WAM; AMS). NEW SOUTH WALES: nr. Fingal; Broken Bay; Palm Beach and Careel Bay, Pittwater (all AMS); Sydney (MCZ, AMS, USNM, MHNP); Botany Bay; Woolaware Bay, nr. Cronulla (both AMS); Wollongong (ANSP); Jervis Bay, Huskisson; Narooma, (both AMS). WESTERN AUSTRALIA: Red Bluff, Shark Bay; Dirk Hartog Island; Herald Bight, N.E. side of Peron Peninsula, Shark Bay; Denham, Peron Peninsula, Shark Bay (all WAM); Carnavon (WAM, USNM); 20 mi. N. of Cardabia; Monte Bellow Islands (both WAM); Barrow Island (WAM, USNM); Cape Bossut (BPBM); mouth of False Cape Creek, La Grange Bay (USNM, ANSP); Broome (AMS, MCZ, USNM, ANSP); Buccaneer Archipelago; Vansittart Bay (both AMS). NORTHERN TERRITORY: Napier Bay, Melville Id. (WAM); Darwin (MCZ, USNM, ANSP). NEW GUINEA: Hollandia; Seleo Island, off Aitape; Milne Bay; Hilimoi Mission, Milne Bay, Papua (all USNM); Collingwood Bay; Woodlark Island (Murua Island) (both AMS); Port Moresby (USNM, AMS, MCZ, BPBM, ANSP); Oriomo River, Papua; Daru, Papua (both MCZ); Merauke (MCZ, USNM); Fakfak (MCZ); 4 mi. N.W. of Korido Village, Soepiori, Schouten Islands (ANSP); Sorendidori Bay, Soepiori, Schouten Islands (MCZ); Samberbaba, Japen Island; Rainbawi Point, Japen Island; '/2 mi. E. of Kaipoeri Village, Koeroedoi Island, INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 461 Geelvink Bay (all ANSP); Mios Woendi, Schouten Islands (USNM): ADMIRALTY ISLANDS: Koruniat Island (ANSP). BISMARKS: New Britain (AMS); New Ireland (MCZ). SOL- OMON ISLANDS: Malaita (MCZ; BPBM); Three Sisters Island; Marava Lagoon, New Georgia (both MCZ); Pavuvu Island, Russell Ids. (USNM); Florida Island (MCZ, USNM); Tulagi, Florida Island (MCZ); Guadalcanal (USNM, AMS). SANTA CRUZ GROUP: Sunday Bay, Vanikoro Island (AMS). NEW HEBRIDES: Qakea, (AMS); Ver, N.E. Santa Maria Id., both Banks Group (USNM); Hog Harbor, Espiritu Santo Island (MCZ); Eromanga (AMS). NEW CALEDONIA: mouth of Ponandou River, E. of Touho (ANSP); Houailoo (BPBM); Ile Nou; Baie du Prony, S. side Mount Dore, nr. Noumea; Ricaudy Reef, Noumea; la Roche Percée, Bourail (all ANSP); llot Amédie (AMS). FIJI ISLANDS: Yasawa Islands (USNM); Savusavu, Vanua Levu (MCZ); N. shore Nananu-i-ra Island, (MCZ, USNM):; Suva (MCZ, USNM, BPBM); Nadronga (USNM, MCZ); Mbalavu (BPBM); Korolevu (ANSP); Lautoka (MCZ); Nadi Airport, all Viti Levu (USNM); Onega Levu; Navutu-i-loma (both BPBM). MARIANAS: Saipan (ANSP); Guam (USNM, BPBM). PALAU ISLANDS: Babelthaup Island (USNM, BPBM, ANSP); Koror Island (BPBM); Malakal Pass, W. of Aurapushokaru Island; S.E. end of Eil Malk (both ANSP); Peleliu (BPBM). CAROLINE ISLANDS: Yap Island, (MCZ, ANSP); Moen Island; Dublon Island, both Truk Is- lands (both BPBM); Ponape (BPBM); Kusaie Island (USNM, BPBM); Kapingamarangi (USNM); MARSHALL ISLANDS: Eniwetok; Arno; Jaluit Atolls (all USNM). GILBERT IS- LANDS; Abaiang Island (BPBM, ZMA); Kingsmill Group (ANSP, USNM). HAWAIIAN ISLANDS: Kauai; Oahu; Molokai and Hawaii Islands (Many localities, see map). LINE ISLANDS: Palmyra Island (MCZ, BPBM); Fanning Island, (BPBM). ELLICE ISLANDS: Vaitupu; Nukulailai (both USNM). WALLIS ISLANDS: btwn, Luaniva and Fungalei Islands (USNM). TONGA: Velitoa Tongatapu (BPBM). SAMOA: Toamua (BPBM); Saluafata, both Upolu (ANSP); Pago Pago (BPBM, USNM, ANSP); Fagaitua Bay, both Tu- tuila(MCZ). SOCIETY ISLANDS: Fanui Bay (USNM); N. end Vaitape; between. Pererau Fai and Araara Pupu, District of Anau, all Bora Bora (both ANSP); Bay Vaiore; around point from Hipu, both Tahaa; Uturoa; Tevaitoa, all Raiatea; Lac Maheva; Port du Bourayne, both Huahine; W. fork of the Opunohu River, Moorea; N.W. of mouth of Nahoata River, Pirae; 9 km. from Taravao, Vairao, Taiarapu Peninsula; Motu Fenuaino, Tautire (all USNM); S. side of Faratea Point, Faaa District, all Tahiti (ANSP). GAMBIER ISLANDS: Rikitea, Mangareva; Vaituatai Bay, Mangareva, (both BPBM). 20 40° 60 80 \oo° 120 140° (60° 180 160° 140° 120° 27 : 3 : s ; | Re PS | aa===--\ [ER NL ee ON Le la i | 20 +— - ANDAMAN ¢ ) wae ! \s NICOBAR), x NSB Me ay CEYLON 5 MALDIVE » ae: : e » ey e GHOND I. coe e SCABRA 2 ise es > MARQUESAS IS. | COLOD KEELING, | ATOLLS TUAMoTY | ARoy gets : a5e : ao ee lee eee __ cateDont a gs eh pea NEW ZEALAND > = . = = 4 n i 4 n 1 a 40 66 80 Too" 120° 140° 160" 180" 160° 140° ; 120" Plate 354. Geographical distribution of Littorina (Littorinop- sis) scabra scabra (Linné) in the Indo-Pacific faunal region. [05-335] 462 _Littorinopsis Littorina melanostoma Gray, 1839 (Pls. 325, 355) Range—Southeast coast of India to Borneo. Remarks—The “Black-mouth” littorine is a very distinctive species within its rather narrow range in south Asia and the East Indies. Its graceful, attenuate outline, dark brown columella Joseph Rosewater Littorinidae and color pattern offer a combination of charac- ters enabling rapid identification and may qualify this species for the title “most attractive Lit- torina.” Nearest relatives appear to be L. cari- nifera Menke, which also has a fairly restricted distribution in the same general region, and L. scabra Linné which is pan-tropical. Among scabra, carinifera and melanostoma, there are Plate 355. Littorina melanostoma Gray (figs. 1-4) Littorina carinifera Menke (figs. 5-13). Figs. 1,2. Lectotype of L. melanostoma Gray, from Pe- nang, Malaysia (BM(NH) 1968364, 23.8 x 11.6 mm.). Figs. 3,4. Port Swettenham, Malaysia (USNM 661028, 28.1 x 13.9 mm.). Fig. 6. Lectotype figure of Phasianella carinifera Menke (from Philippi, Abbildungen and Beschreibungen Con- chylien, vol. 2, pl. 5, fig. 22; about 21 x 12 mm.). Figs. 5,7. Sandakan, North Borneo; note similarities to lectotype figure (USNM 658105a, 16.5 x 10.9 mm.). Figs. 8,9. Lectotype of Littorina rubropicta von Martens, from King Id. Bay, Mergui Archipelago, Burma (BM(NH) 87.3.10.140-144, 18.3 x 10.5 mm.). Figs. 10,11. Lectotype of Littorina conica Philippi, from Java (BM(NH) 1968225, 22.4 x 12.7 mm.). Figs. 12,13. Bombay, India (USNM 90470, 21.6 x 12.4 mm.). [05-336] November 30, 1970 many similarities. All three occupy a_ similar habitat, anatomical details are remarkably simi- lar, and although spawning behavior has not been verified for melanostoma and carinifera, they very probably reproduce ovoviviparously. Habitat—Estuarine areas on mangrove trees and grasses. Description—Shell reaching 28.2 mm. (about 1 inch) in length, attenuately conical in shape; average obesity about .48 (23 specimens ranged from .44 to .54); relatively thin but strong in structure, imperforate, and sculptured with shal- lowly incised spiral striae, overall microscopic wavy spiral threads and fine, oblique axial lines of growth. External coloration rather uniform; ground color yellowish white, with closely spaced punctate to elongate-rhomboidal brown markings located between the spiral striae, usual- ly arranged axially but sometimes obliquely or in zigzag pattern; sometimes with white dashes be- tween brown spots; tip of spire often gray; aper- ture yellowish white or with outer brown color markings partially showing through; columella dark chocolate-brown. Base hardly flattened; only a very weak keel at periphery. Whorls 6-8, usually rather flat-sided. Spire usually somewhat more than half the length of shell, produced at an angle of 42-44°. Aperture oval; outer lip rather thin, often narrowly shouldered; inner lip not greatly thickened, weakly concave, the posterior 2/3 glazed with dark brown callus, white where it joins outer lip at anterior extremity. Suture weak- ly impressed. Sculpture consisting of about 6 shallow spiral striae on spire whorls, persisting onto body whorl where as many as 18 may be present over length of whorl; area between striae flattened and each may be divided by single, weak, secondary stria; entire surface covered by fine microscopic wavy spiral threads; floor of striae often appear to be finely, shallowly and closely punctured. Axial sculpture consisting of rather regularly spaced oblique growth lines be- coming coarser near outer lip. Operculum large, corneous and paucispiral. Periostracum not evi- dent in specimens examined. Nuclear whorls par- tially decollate in all specimens examined; re- maining portions smooth and glassy, light grayish tan in color; succeeding whorls sculptured spiral- ly. Radula typically littorinid (2-1-1-1-2); central tooth broad and low (similar to L. scabra and L. carinifera). Animal darkly pigmented on surfaces of ten- tacles snout and foot. External appearance of animal removed from shell: in female the “‘cover- INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 463 ing-capsule gland complex” (see Fretter and Graham, 1962) located at posterior right of cteni- dium, is very distinct, pigmented, and forms a spiral pattern; in male verge is greatly thickened from base to one-half its length; distal portion filamentous; sperm duct deeply folded on floor of mantle cavity and along posterior edge of thick- ened base of verge and inner edge of filamentous portion. Nothing reported concerning spawning or development of this species, although it is suspected that species is ovoviviparous. In both male and female the ctenidia are greatly enlarged and occupy most of the area of mantle cavity roof, darkly pigmented and conspicuous. Measurements (mm.)— length width no. whorls locality 28.2 14.1 8 Port Swettenham, Malaysia 27.8 14.1 8 Chaga, Malaysia 5.1 12 8 Nakhorn Si Thammarat, Thailand 25.9 12.9 7+ Nakhorn Si Thammarat, Thailand 24.4 11.5 7 Ko Chang, Thailand 24.0 10.9 7 Port Swettenham, Malaysia 22.6 11.1 T+ Sarawak, Borneo 22.3 12.0 7 Kranji, Singapore 21.9 9.6 8 Burma 21.1 10.1 8 Laem Ngob, Thailand 20.4 102 6+ Port Canning, India 20.0 9.7 8 Vietnam 19.2 9.4 6+ Taiwan 14.9 ie 5+ Hong Kong J Ke _[MELANOSTOmaA] *% COCO - KEELING ATOLLS Pe 0° 80° 100° 120° \40° Plate 356. Geographical distribution of Littorina (Littorinop- sis melanostoma (Gray) in south Asia and Indonesia. [05-337] 464 Littorinopsis Synonymy— 1839 Littorina melanostoma Gray, The Zoology of Captain Beechey’s Voyage—in His Majesty's Ship Blossom, Mollusks, pl. 140 (Indian Ocean; [Penang, Malaysia, here restricted]) lectotvpe B.M. (NH) 1968364: 23.8 x 11.6 mm.; non L. melanostoma A. J. Krynicki, 1837, Bull. Soc. Imp. des Nat. de Moscow, Ann. 1837, No. 2, p. 60, nomen nudum. 1885 Littorina melanostoma articulata Nevill, Hand List of Mollusca in the Indian Museum, part 2, p. 151 (Hong Kong; type in Indian Museum, Calcutta). Types—Littorina melanostoma apparently is one of the species not collected during the voyage of the Blossom as Gray gave for a type locality, “Indian Ocean” which the expedition did not reach (Rosewater, 1968). One of the 5 syntypes from Gray’s collection is here selected as lecto- type: BM(NH) 1968364, 23.8 x 11.6 mm (pl. 355, figs. 1, 2). The type locality is here restricted to Penang, Malaysia. Records—INDIA: Port Canning (USNM, RNHL); Manali, off Mandapam (MCZ). CEYLON: (E. von Martens, 1887). BURMA: King Island Bay, Mergui Ids. (E. von Martens, 1887). THAILAND: Ko Chang (USNM); Ang Hin (MCZ); Ban Tamru, both Cholburi Province (ZMA); Laem Ngob; Pak Phun (both USNM); Pak Paying, Nakhorn Si Thammarat (MCZ). MALAYSIA: N.E. corner of Pulau Lumut, Port Swet- tenham (USNM); Malacca (MCZ). SINGAPORE: Kranji (ANSP, USNM). CHINA: Hainan; Tsi Mei, Amoy; Hai- ch’eng, Fukien Province (all ANSP); Castle Peak Bay, Hong Kong (NMW). TAIWAN: (USNM). VIETNAM: Saigon River, Cochin China (USNM, MCZ). INDONESIA: Palau Weh (RNHL); Telok Niboeng (ZMA); Tandjung Tiram, all Sumatra (ZMA); Djepara (RNHL); Surabaja, both Java (ZMA; RNHL; SMF); Santubong, Sarawak, Borneo (MCZ, USNM). Littorina carinifera (Menke, 1830) (Pls. 325, 355) Range—South and southeast Asia from West Pakistan to the Philippines and Borneo. Remarks—Of the several forms which have been described as being closely related to Lit- torina scabra, only two appear worthy of recogni- tion as distinct species: Littorina melanostoma Gray and Littorina carinifera Menke. The latter, “carinate littorine,’ in its most extreme form, lives up to its descriptive name often having as many as 3-5 rather strong carinae on the body whorl. Variation is considerable, however, and there are populations in which carinae are present only at the periphery. The species may be recog- nized by its rather pyramidal shape, the color pattern of rather straight to oblique or zigzag reddish axial bands on a yellowish brown back- ground, and by the presence, usually, of dark color bands within the aperture. Joseph Rosewater Littorinidae Habitat—On bushes or low shrubs along shore; mangrove swamps. Description—Shell reaching 22.2 mm. (about 0.9 inch) in length, pyramido-conical in shape; average obesity about .63 (23 specimens range from .55 to .71); relatively thick in structure, imperforate, often multiply carinate; otherwise sculptured with well-marked spiral striae, overall microscopic wavy spiral threads (often obscured by wear) and fine, regular, closely spaced axial lines of growth. External coloration rather uni- form; ground color yellowish to light-brown, with axial pattern of nearly straight to oblique reddish brown lines. In carinate specimens pat- tern consists of reddish brown spots primarily on carinae. Aperture grayish white with three or more dark brown bands revolving within; colu- mella white. Base somewhat flattened; periphery, at least, markedly carinate. Whorls 6-7, hardly rounded—to rather flat sided. Spire usually some- what more than half the length of shell, produced at an angle of 46-72° (depending on specimen). Aperture roundly oval; outer lip rather thin but sometimes thickened within; inner lip not greatly thickened, occasionally with a denticulate swel- ling near its base, white in color. Suture moder- ately to well impressed in highly carinate speci- mens. Sculpture consisting of about 10 often punctate spiral striae on spire whorls, persisting onto body whorl where as many as 20 may be present over length of entire whorl; in non- strongly carinate specimens areas between striae flattened to moderately well rounded as incipient carinae; in strongly carinate specimens at least 3 inter-stria areas raised as rather strong carinae and others may be moderately raised; in these speci- mens striae are deep furrows and occasionally may be sculptured intrinsically with closely- spaced axial wrinkles; entire surface covered by fine microscopic wavy spiral threads, the latter often obliterated by wear. Axial sculpture consist- ing of rather regularly-spaced axial growth lines. Operculum moderate in size, corneous, pauci- spiral. Periostracum not evident in specimens examined. Nuclear whorls partially decollate in all specimens examined; remaining portions smooth, shining, light tan in color; succeeding whorls spirally sculptured. Animal including radula resembling L. scabra and L. melanostoma; darkly pigmented on upper surfaces of tentacles, snout and foot. Ctenidial area darkly pigmented showing through mantle; in female “covering-capsule gland complex” (see Fretter and Graham, 1962) located at posterior [05-338] November 30, 1970 right of ctenidium is very distinct, its spiral outline marked with dark pigment; in male verge greatly thickened at base and for about one-half its length; thickened portion terminally bulbous; distal portion filamentous; sperm duct deeply folded, running along posterior edge of verge. Nothing reported concerning spawning or devel- opment, but probably ovoviviparous. Measurements (mm.)— length width no. whorls locality 29:9 1252 7 Bombay, India 19.0 112 7 Manapla, Negros Occi- dental, Philippines 18.6 12 6 Zamboanga, Mindanao, Philippines 17.8 11.1 7 Iloilo, Panay, Philippines 17.0 10.6 7 Singapore 16.7 11.4 6 Pancol, Palawan, Philippines 16.4 11.2 6 Po Bui Id., Sandakan, North Borneo 14.3 9.0 7 Mangalore, India 13.8 8.4 6+ Silaga R., Samar, Philippines 10.4 74 4+ Djakarta, Java Synonymy— 1830 Phasianella carinifera Menke, Synopsis methodica Mol- luscorum, Pyrmont, edition 2, pp. 51, 141 (locality unknown). 1832 Littorina perdix King and Broderip, Zoological Journal, vol. 5 (1831), p. 345 (no locality given), 1846 Littorina conica Philippi, Proceedings of the Zoological Society of London, part 13 (1845), p. 141 (Java); lectotype BM(NH) 1968225: 22.4 x 12.7 mm.; 1847, Abbildungen und Beschreibungen Conchylien, vol. 3, p. 9, Litorina, pl. 6, figs. 1, 2. 1885 Littorina carinifera pyramidalis Nevill, Hand-List Mol- lusca in Indian Museum, part 2, p. 151 [nomen nodum |]. 1885 Littorina carinifera laevior Nevill, ibid. [nomen nudum ]. 1887 Littorina rubropicta von Martens, Journal of The Lin- nean Society, Zoology, vol. 21, p. 170, pl. 16, figs 2 a-f (King Island Bay, Mergui Archipelago); lectotype BM(NH) 87.3.10. 140-144: 18.3 x 10.5 mm. 1932 Litorina carnifera Sherborn, Index Animalium, section 2, part 31, p. 638 [error for L. carinifera]. Types—According to Dance (1966) Menke’s collection was dispersed and therefore, it is most unlikely that the original type specimen of L. carinifera can be located. Fortunately Philippi (1847, Abbildungen und Beschreibungen Con- chylien, vol. 2, Litorina, p. 227, pl. 5, fig 22) apparently had access to Menke’s collection and INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 465 MALDIVE » * 1s "¢ ele _ ~s = is * a NEW CHAGOS | ~ 5 & Suir isco 7 a ES COCOG - KEELING ATOLLS ‘e CARINIFERA “Mouritias t — — — ee ee ee A ct Aare wees =a! n ale o° 80° 100° : yzo° (40° Plate 357. Geographical distribution of Littorina (Littorinop- sis) carinifera (Menke) in South Asia, Indonesia and the Philippines. figured what he called “the original example”. That figure is here selected as the lectotype of Phasianella carinifera Menke (pl. 355, fig. 6). The type locality is here designated as Negros Oc- cidental, Philippines, one of the localities men- tioned by Philippi. The location of the type-specimen of L. perdix King and Broderip is unknown. It may be in the BM(NH) although it was not found during a recent visit. The lectotypes of L. conica Philippi and L. rubropicta von Martens are in the BM(NH) as mentioned in the synonymy (pl. 355 figs. 8-11). Records—WEST PAKISTAN: China Creek, Karachi (MCZ). INDIA: Bombay (MCZ, USNM); Vengurla, N. of Goa; Netravati River, Mangalore (both USNM). BURMA: King Island Bay, Mergui Ids. (E yon Martens, 1887). THAILAND: Ko Kut (USNM). MALAYA: N.E. corner Pulau Lumut, Port Swettenham; Singapore (both USNM). PHILIPPINES: Medio Island, Galera Bay, Mindoro; Hoilo, Panay; Victorias; Manapla, both Negros Occidental; W. of Catbalogan; Silaga River, both Samar; Zamboanga, Mindanao; Pancol, Palawan (all USNM); Busuanga (RNHL). INDONESIA: Belawan, Su- matra (RNHL; ZMA); Panahatan, Sumatra; Karanganjar, Java (both ZMA); Tandjung Priok, Java (RNHL); Djakarta, Java (ZMA); Santubong, Sarawak, Borneo (both USNM); Labuan, Borneo (NMW, RNHL); Sipitang, North Borneo; Po Bui Island, Sandakan, North Borneo (both USNM). [05-339] 466 Littorinopsis ?Littorina miodelicatula (Oyama, 1950) (Pl. 358, figs. 1-3) Range—Tertiary of Japan. Remarks—The_ species was originally de- scribed as resembling some of the Trochidae but Oyama preferred an assignment in Littorinidae near L. scabra because of residual color pattern in the fossil and a shape somewhat like that of L. carinifera. It is included here somewhat doubt- fully as the affinity with Archaeogastropoda seems most likely. Synonymy— 1950 Littorinopsis (Littorinopsis) miodelicatula Oyama, Geo- logical Survey of Japan, Report no. 132, p. 8, pl. 1, figs 2, 3. (Ueno, Japan; Tertiary Kakebata formation) holotype in collection of Geological Survey of Japan [?]: 8 x 6.7 mm. Littorina incisa Yokoyama, 1927 (Pl. 358 figs. 4, 5) Range—Pliocene of Japan. Remarks—Littorina incisa is a relatively small but well-preserved fossil. The excellent illustra- tion given by Yokoyama and reproduced here indicates that this species is probably most close- ly related to L. melanostoma and is quite unlike any other Recent species because of its attenuate shape. Synonymy— 1927 Littorina incisa Yokoyama, Journal of the Faculty of Science, Imperial University of Tokyo, section II, Geology, Mineralogy, Geography, Seismology, vol. 2, part 4, p. 175, pl. 47, fig 8 (Pliocene, Nagaya, Kaga, Japan); holotype in Geological Institute, Imperial University of Tokyo: 5 x 2.5 mm. Joseph Rosewater Littorinidae Plate 358. Figs. 1-3. Littorinopsis miodelicatula Oyama, from Tertiary, Ueno, Japan (Holotype, from original figures in Report no. 132, Geological Survey of Japan, pl. 1, figs. 3a-c; 8 x 6.7 mm.). Figs. 4,5. Littorina incisa Yokoyama, from Pliocene, Nagaya, Kaga, Japan (Holotype from original figures; 5 x 2.5 mm.). [05-340] November 30, 1970 Subgenus Austrolittorina new subgenus Rosewater Type-species: Littorina unifasciata unifasciata Gray, 1826 The members of this subgenus are character- ized by a predominantly southern ocean and tropical distribution and by the possession of shell and anatomical features generally similar to those of its type-species, L. unifasciata unifas- ciata: a conico-turbinate shell, flattened columel- la and crescent-shaped area on the adjacent part of the shell; verge with a basal enlargement incorporating a single penial gland containing an accessory flagellum. Littorina unifasciata Gray, 1826 The Littorina unifasciata species complex is represented in the Indo-Pacific faunal region by three entities, the nominate subspecies, unifas- ciata in Australia, and the subspecies antipoda in New Zealand and fernandezensis in the Juan Fernandex Islands. All closely resemble each other except for size or proportional differences which are the result, probably, of isolation. All members of the subgenus Austrolittorina show a preference for the southern ocean south of the equator. Littorina unifasciata subspecies unifasciata Gray, 1826 (Pls. 325, 326, 359-361) Range—Southern coast of Australia, predomi- nantly south of the Tropic of Capricorn, from Carnarvon in the west to Queensland; Lord Howe Island; Tasmania. Remarks—The nominate subspecies of the L. unifasciata group is distinctly a native of the Australian continent occurring extraterritorially only in Tasmania and on Lord Howe Island. The plot of its distribution, if smoothed, would resem- ble an inverse normal curve (pl. 362); it is re- stricted almost exclusively to the more southern coasts south of the Tropic of Capricorn. This species has been confused in the literature with L. mauritiana Lamarck, although the two are quite distinct morphologically and their ge- ographic ranges are separate (see mauritiana). INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 467 Both Reeve (1858) and Tryon (1887) considered unifasciata to be a synonym of mauritiana. Al- though both may have the single dark band around the whorls upon which the name “unifa- sciata” is based, mauritiana is often larger and more elongate, it bears subtle darker patterning and lacks the columellar crescent and _ penial gland of unifasciata. Habitat—Lives on shore rocks in the spray zone, above high water. Description—Shell reaching 20.9 mm. (about .8 inch) in length, conical to subturbinate in shape, average obesity about .62 (45 specimens range from .55 - .70); older individuals moderately thick in structure, imperforate, usually developing a flattened, crescent-shaped area adjacent to the columellar callus; sculptured with spiral striae occasionally reinforced with fine brown color lines, spiral, white microscopical textural threads, and irregular, oblique, axial lines of growth. Ex- ternal ground color grayish white to bluish gray, usually the former with a rather diffuse blue-gray band encircling the body whorl and anterior portion of spire whorls; apex light brown. Aper- ture medium to dark-brown with a prominent white band inside near junction of outer lip and columella. Base somewhat flattened, separated from upper part of body whorl by a low but distinct keel at periphery. Whorls 5-7, rather straight-sided. Spire less than half the length of shell, convex, produced at an angle of about 60°. Aperture oval to subquadrate; outer lip moder- ately thick, having its origin high on body whorl above keel so that keel enters aperture; outer lip striated internally at edge, reflecting external sculpture; inner lip weakly concave. Columella dark brown to white, shallowly excavated, some- what rimmed medially, and with a flattened brown to white crescent-shaped area distally on the adjacent base. Suture not deeply impressed. Sculpture consisting of from 6-9 spiral striae on spire whorls, persisting onto surface of body whorl where as many as 12 may be present above the keel; becoming indistinct below keel. Entire surface covered with microscopic, closely spaced rather straight spiral white threads partially im- bedded in shell substance and seen through the surface; not detectable in worn specimens. Axial sculpture consisting of rather irregular oblique lines of growth. Operculum corneous, moderately thick, paucispiral. Periostracum not evident in specimens examined. Nuclear whorls decollate in most mature specimens; when present, light brown in color, smooth and about 3 in number; [05-351] 468 Austrolittorina first post-nuclear whorl similarly colored, but sculptured with spiral striae. Radula littorinid (2-1-1-1-2) central tooth somewhat narrow. Animal darkly pigmented on surfaces of ten- tacles, snout and foot. Verge moderately short and thick, yellowish white in color; having a basal Joseph Rosewater Littorinidae flap with a single penial gland containing an internal hyaline accessory flagellum. Nothing is known concerning the reproduction and develop- ment of this species, although probably it is oviparous and spawns pelagic capsules. Plate 359. Subspecies of Littorina (Austrolittorina) unifasciata (Gray). Figs. 1,2. L. unifasciata unifasciata from Port Jackson, New South Wales, Australia (USNM 89472; 20.9 x 12 mm.; note flattened crescent sloped area adjacent to colu- mellar callous). Figs. 3,4. L. unifasciata unifasciata from Red Bluff, Kalbar- ri, Western Australia (USNM 691673; 12.7 x 7.6 mm.). Fig. 5. L. unifasciata unifasciata from same locality as Figs. 3,4; detail of shell surface to show white microscop- ical textural threads (enlargement of shell area measuring about 1.3 x .8 mm.; young specimen, length 4.9 x 3.2 mim.). Figs. 6-8. L. unifasciata antipodum; fig. 6, from Island Bay, Cook Strait, New Zealand (USNM 671202, 8.7 x 4.8 mm.); figs. 7,8, from Taipa, Doubtless Bay, North Island, New Zealand (USNM 601974; 7.4 x 4.7 mm.). Figs. 9-12. L. unifasciata fernandezensis from eastern shore Cumberland Bay, Isla Mas a Tierra figs. 9, 10, Paratype (USNM 679256, 11.8 x 7.2 mm.); figs. 11, 12, Holotype (USNM 368900, 13.4 x 7.8 mm.), [05-352] November 30, 1970 Measurements (mm.)— length width no. whorls locality 20.9 11.9 5+ Port Jackson, New South Wales 19.9 12.9 5+ Port Jackson, New South Wales 19.3 11.3 5 Port Jackson, New South Wales 18.2 11.4 6 Kalbarri, Western Australia 17.9 11.6 4+ Kalbarri, Western Australia 16.3 9.5 5+ Robe, S. E. Australia 16.0 9.6 5 Port Denison, Western Australia 15.3 9.3 5+ Port Fairy, Victoria 14.3 9.1 5 Mouth of Murchison River, Western Australia 13.0 7.9 4 near Sydney, New South Wales 1:29 8.1 4 Mouth of Murchison River, Western Australia 10.9 6.5 4+ Queenscliff, Victoria 10.7 Wee 5 Wollongong, N.S.W. 9.4 6.1 4 Coogie Beach, Sydney, N.S.W. Heo 4.7 5 Coogie Beach, Sydney, N.S.W. 6.4 3.8 3+ Coogie Beach, Sydney, N.S.W. 5.0 Sill 3+ Pearl Beach, N.S.W. 4.9 3.4 3+ between Port Phillip Heads and Cape Otway, Victoria Synonymy— 1826 Littorina unifasciata Gray, in P. P. King, Narrative of a survey of the coasis of Australia, vol. 2, Appendix B, p. 483 (Australia [King George Sound, Western Aus- tralia, here restricted ]); lectotype B.M.(N.H.) 1968373: 16.4 x 10.6 mm. 1833 Littorina diemenensis Quoy and Gaimard, Voyage de decouvertes de ]’Astrolabe, vol. 2, part 2, p. 479, pl. 33, figs. 8-11 (rocks of the littoral zone of South Australia, Tasmania and also New Zealand [here restricted to Tasmania]); lectotype in Mus. d Hist. Nat., Paris: 10.2 x 5.9 mm. 1843 Litorina acuta Menke, Molluscorum Novae Hollandiae Specimen, p. 9 (Western Australia; type lost); 1844, Zeitschrift fiir Malakozoologie, Jahr, 1844. p. 57. 1847 Litorina mauritiana crassior Philippi, Abbildungen und Beschreibungen Conchylien, vol. 2, p. 165, Litorina pl. 3, fig 17[a] (Australia [here restricted]). 1850 Littorina diemensis Gray, Figures of Molluscous Ani- mals, vol. 4, p. 78 [error for L. diemenensis Quoy and Gaimard, 1833]. 1858 Littorina laevis ‘Philippi’ Reeve, Conchologia Iconica, vol. 10, Littorina, pl. 17, fig 95 (locality not given); not L. laevis Philippi = L. mauritiana Lamarck. 1885 Littorina diemenensis pseudolaevis Nevill, Hand-List of Mollusca in the Indian Museum, Calcutta, part 2, p. 141, (New South Wales); new name for L. laevis Reeve, not L. laevis Philippi. Types—The lectotype of L. unifasciata Gray is in the BM(NH), 1968373. The type locality men- tioned by Gray, Australia, is here restricted to INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 469 Plate 360. Figs. 1,2. Lectotype of Littorina unifasciata Gray from Australia [King George Sound, Western Australia] (B. M.(NH) 1968373, 16.4 x 10.6 mm.). Figs. 3,4. Lectotype of Littorina diemenensis Quoy and Gaimard [= L. unifasciata] [from Tasmania] (MHNP, 10.2 x 5.9 mm.). Fig. 5. Lectotype figure of Litorina antipodum Philippi, from Abbildungen und Beschreibungen Conchylien, vol. 2, p. 195, pl. 4, fig. 2 (New Zealand, about 7.8 x 4.5 mm.). King George Sound, Western Australia, one of the localities visited during King’s survey. The lecto- type of L. diemenensis Quoy and Gaimard is in the Paris Museum. It is suspected that the illus- tration is a composite since there were no speci- mens in the type lot which fully resembled it. A lectotype was chosen which most closely approxi- mated the illustration, although smaller than the original measurements (10.2 x 5.9 mm. versus approximately 11.3 x 6.8 mm. (5x 3 lignes)). The holotype of Menke’s L. acuta may be presumed lost; however the author himself (1844, see Syn- onymy) synonymized his species with unifasc- iata an action which is accepted here as proof of the identity of the species in view of the absence of a type specimen. The type of L. crassior Philippi may be in the Berlin Museum; the type of L. pseudolaevis Nevill may be in the Indian Muse- um, Calcutta. Records—AUSTRALIA: QUEENSLAND: Point Lookout, Stradbroke Island, Moreton Bay (WAM, AMS); Freshwater; Noosa Heads; Caloundra; Coolangatta (all AMS). NEW SOUTH WALES: Byron Bay (MCZ, AMS); Yamba (MCZ); Merewether Beach, Newcastle (WAM); Pearl Beach: The [05-353] 470 Austrolittorina AI AP YT ; 5mm Plate 361. Fig. A. Radula of Littorina unifasciata unifasciata from The Entrance, New South Wales, Australia, (USNM 631284; upper scale is .05 mm.) Fig. B. Radula of L. unifasciata fernandezensis from Mas Afuera Id., Juan Fernandez Islands (middle scale is .1 mm.). Fig. C. Penis of L. unifasciata unifasciata from Kalbarri, Western Australia, USNM 691673 (lower scale is 5 mm.). Entrance (both USNM); Putty Beach (AMS): S. shore, Broken Bay, nr. entrance (ANSP); btwn. Newport and Palm Beach (ANSP, USNM; Middle Harbor (AMS); Long Reef, N. of Manly, nr. Sydney (AMS, ANSP); Sydney (MCZ): Port Jack- son (USNM, MCZ, AMS, ANSP); La Perouse, Botany Bay (MCZ, ANSP, WAM); Kurnell (AMS); Wollongong (AMS, USNM); Port Kembla (USNM); Merimbula; Two Fold Bay (both MCZ, AMS), VICTORIA: Mallacoota (AMS); Flinders (USNM); Sandringham, Port Philip (ANSP, USNM):; Port Philip (ANSP, AMS, MCZ); btwn. Port Philip Heads and Cape Otway (USNM); Barwon Heads (MCZ); Port Fairy (ANSP, USNM, AMS, NMW); Bridgewater Bay (MCZ); Port- land (MCZ); TASMANIA: Fischer Island, Bass Strait (AMS); Islets S. of Flinders Island, Bass Strait (WAM); King Island; Bicheno; Northwest Long Bay, Port Arthur; Simpson’s Bay, D’Entrecasteaux Channel (all AMS); Eagle Hawk Neck; Ho- bart (both MCZ); Taroona, Derwent River; Grant’s Head, Sloop Rocks (both AMS); Low Head (MCZ). SOUTH AUS- TRALIA: Boatswain Point (Cape Thomas) nr. Robe (ANSP, USNM, MCZ); Robe (NMW); Encounter Bay; Port Willunga (both MCZ); Adelaide (USNM); Giles Point, Yorke Peninsula (USNM, MCZ, NMW); Wool Bay (NMW); South Neptune Island, Spencer Gulf (MCZ, AMS). WESTERN AUSTRALIA: Mondrain Island, Recherche Archipelago (WAM); Esperance (AMS); mouth of Pallinup R. Estuary (WAM); Frenchman’s Bay, nr. Albany (AMS); Oyster Harbor, Nr. Albany, N. of Bayonet Head; Middleton Bay, nr. Albany; Windy Harbor, Nr. Northcliffe (all WAM); Cape Leeuwin, (AMS); Margaret River (WAM); Wallcliffe, nr. Margaret River (MCZ); Yallingup, Cape Naturaliste; Bunker Bay, Cape Naturaliste (both USNM); Geographe Bay; Bunbury (both AMS); Cottesloe (WAM, AMS): Point Peron, nr. Perth, on boulders; Rottnest Island (both MCZ, AMS); Perth (AMS); Little Anchorage, Leeman; Snag Island, Leeman; Beagle Islands (all WAM); Port Denison (USNM); Greenough River mouth; Harrocks Beach nr. Northampton (both WAM): Red Bluff, Kalbarri, nr. mouth of the Murchison River (WAM, USNM); Eagle Bluff, Freycinet Estuary (WAM); Point Gregory, N.W. Corner Peron Peninsula (WAM); Dirk Hartog Island, (last three all Shark Bay) (AMS); below Quodd’s Light, N. of Carnarvon (WAM), LORD HOWE ISLAND (ANSP, AMS). Joseph Rosewater Littorinidae eT cee a) Poy : : & 2 PHOEN is “ex: » TOK rig AMon ts Q HEORIDES sa Oe etn. | + : mM Ne “Tousa Seen eee my CALEDONIA. “9 a Norfolk Plate 362. Geographical distribution of Littorina (Austrolit- torina) unifasciata unifasciata (Gray), in Australia, and of the subspecies, L. unifasciata antipodum (Philippi), in New Zea- land. Littorina unifasciata subspecies antipodum (Philippi, 1847) (Pl. 359, figs. 6-8, pl. 360) Range—Islands of New Zealand. Remarks—The New Zealand species best known as Littorina oliveri (Finlay) is without a doubt L. antipodum Philippi, here considered a subspecies of L. unifasciata. Finlay’s (1930) com- parison of antipodum with the typical Australian unifasciata best summarizes the differences: “The New Zealand species differs in its smaller shell, higher and more slender spire, with almost straight instead of distinctly convex whorls, no- tably smaller aperture, and darker more promi- nent and better defined blue band .. .” It may be theorized that elements of unifasciata were car- ried to New Zealand in the past and that the subspecific differences evolved in response to the new environmental conditions during isolation. Habitat—On rocks at and above high water mark. Description—Shell reaching 12.3 mm. (about .5 inch) in length; in general appearance looking much like a diminutive L. unifasciata unifas- ciata, elongate to rather short-conical in shape; average obesity about .59 (21 specimens range from .52-.64). A flattened, crescent-shaped area adjacent to columella callus usually present, but often reduced in size or may be lacking. Sculpture similar to L. unifasciata unifasciata. Color band encircling body whorl and anterior portion of [05-354] November 30, 1970 Plate 363. Geographical distribution of Littorina (Austrolit- torina) unifasciata fernandezensis Rosewater in the Juan Fer- nandez Islands, southeastern Pacific. spire whorls very dark bluish-gray, shell often tinted brown above and below the band. Whorls 4-6, apex usually severely eroded. Radula as in L. unifasciata, the central tooth rather narrow. Animal darkly pigmented on surfaces of ten- tacles, snout and foot. Verge similar to L. cincta and L. unifasciata unifasciata: rather short and thick and having a basal mitten-shaped flap which bears a penial gland with an internal hyaline accessory flagellum. Sexually mature males very small. Nothing known concerning reproduction and development, although is prob- ably oviparous and spawns pelagic capsules. Measurements (mm.)— length width no. whorls locality are 6.9 6+ Wellington, New Zealand 123 6.4 54 Wellington, New Zealand 10.3 5.6 6+ Port Waikato, New Zealand 10,2 6.0 6 Wellington, New Zealand 9.7 6.0 44 Christ Church, New Zealand 9.5 6.1 44 Doubtless Bay, North Island, New Zealand 8.8 5.0 44 Island Bay, Cook Str. New Zealand 8.5 ee 5 Napier, New Zealand 7.9 5.0 6 “New Zealand” 5.4 3.2, 5 Auckland, New Zealand Synonymy— 1847 Litorina antipodum Philippi, Abbildungen und Besch- reibungen Conchylien, vol. 2, p. 195, Litorina pl. 4, fig 2 (New Zealand) INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 471 1930 Melarhaphe oliveri Finlay, Transactions of the New Zealand Institute, vol. 61, p. 224, (Hampden, East Coast of South Island, New Zealand; Holotype in Finlay Collection, Auckland Museum); refers to Suter, 1913, p. 188, and to C. E. R. Bucknill, 1924, “Sea Shells of New Zealand,” p. 37, pl. 7, no. 2); not Melarhaphe zelandiae Finlay, 1926, which = L. cincta Quoy and Gaimard. Types—The type of L. antipodum Philippi may be in the Berlin Museum. The lectotype figure is that of Philippi, Abbildungen und_ Beschrei- bungen Conchylien, vol. 2, Litorina, pl. 4, fig 2 (see pl. 360 fig. 5). The holotype of M. oliveri Finlay is in the Finlay Collection, Auckland Museum. It should be pointed out here that Finlay (1926) by error described Melarhaphe ze- landiae intending the name for the New Zealand relative of L. unifasciata. However, the type of zelandiae proved to be identical with L. cincta, another New Zealand species and it was for this reason that Finlay (1930) thought it necessary to describe M. oliveri not realizing that Philippi had already described the species over 80 years be- fore. Records—NEW ZEALAND: NORTH ISLAND: Tauranga (BPBM); Napier; Plimmerton, Wellington; Titahi Beach, in shell sand (all USNM); Island Bay, Cook Strait (AMS, USNM); Worser Bay, Wellington Harbour (MCZ, USNM); New Plymouth (AMS); Taipa, Doubtless Bay (USNM); Rus- sell, Bay of Islands (MCZ); Hen Island; N.W. point on Motutapu Island (both ANSP); Auckland (AMS, ANSP); W. side Wade River, about 15 mi. N.W. of Auckland (USNM, ANSP); Rangitoto (AMS, ANSP); S. coast, Rangitoto Island (USNM); Manukau Harbor; Port Waikato (both ANSP, USNM). SOUTH ISLAND: Picton, Lyttelton (AMS); Taylor's Mistake, Christchurch (USNM); Porto Bello; Dunedin; Gray- mouth (all AMS); Stewart Island (ANSP); Chatham Islands (Suter, 1913). Littorina unifasciata new subspecies fernandezensis Rosewater (Pls. 359, 361) Range—Juan Fernandez Islands, southeastern Pacific. Remarks—Odhner (1922) reported Littorina mauritiana Lamarck from Mas Afuera, Juan Fer- nandez Islands, and referred to its distribution as extending from Mauritius to New Zealand. In the present study it has been found that L. mauri- tiana is a species restricted to the western Indian Ocean, and that L. unifasciata unifasciata and its subspecies L. u. antipodum, inhabiting Australia and New Zealand respectively, are quite distinct. The third subspecies of the L. unifasciata group, named here, is located geographically quite far from its nearest relative. According to Odhner’s [05-355 ] 472 Austrolittorina Joseph Rosewater Littorinidae (ibid.) analysis the faunal affinities of Juan Fer- nandez show a closer relationship to South Amer- ica than to the western Pacific. Further, he found high endemism in the mollusks, indicating long isolation. Nevertheless the Littorina from Juan Fernandez is clearly related to L. unifasciata, but with sufficient differences to suggest that at least subspeciation has occurred. The answer to the question of how L. unifas- ciata and its subspecies became distributed in such widely separated localities will probably never be answered satisfactorily. Although I know of no fossil record, the history of the ancestors of this group probably dates well back into geologic time when land masses and ocean currents are known to have been quite different. It is possible that the group evidences a relict and disjunct distribution from the time when the Antarctic continent was both larger and warmer than it is today. The subspecies living in southern Australia, New Zealand and Juan Fernandez may have evolved from a once more widely dis- tributed species which migrated northward in response to the cooling off of the southern con- tinent. It is also possible that they represent a modern distribution brought about by the pre- dominately western current of the southern ocean. (See discussion of a somewhat similar circumpolar distribution by Abbott, 1968, pp. 183-188). Littorina unifasciata fernandezensis is very similar to the nominate subspecies, differing in being on the average slightly more slender, al- though it is not so slender as L. u. antipodum (see average obesities in Descriptions of L. unifas- ciata and antipodum). There is a superficial simi- larity in appearance between L. fernandezensis and L. paytensis Philippi of western South Amer- ica, but the former lacks the brown color mark- ings, two white bands in the aperture and single strong subsutural stria which are all present in paytensis. The two also are members of different subgenera of Littorina! Habitat—Shore rocks. Description—Shell reaching nearly 18 mm. (about .7 inch) in length, conical to subturbinate in shape average obesity about .60 (52 specimens ranged from .55 - .64); appearance very similar to L. unifasciata unifasciata, but more turbinate in shape and less conical; columella not as strongly developed; adjacent crescent-shaped, flattened area persists and may be well developed in some specimens. Spiral striae often strongly impressed, 16-22 striae on body whorl above periphery (14- 16 in L. unifasciata unifasciata); striae not rein- forced by brown color lines; spiral sculpture sometimes apparent inside edge of outer lip as a series of fine teeth. Color of aperture varying from brown to violet; color band on body whorl often narrower and darker than in unifasciata. Animal characters similar to L. unifasciata unifasciata Measurements (mm.)— length width no. whorls locality 17.6 11.3 34 Santa Clara Id. Islas Juan Fernandez 16.3 10.2 5+ Cumberland Bay, Isla Mas a Tierra (paratype) 15.7 9.0 5+ Cumberland Bay 15.3 9.9 4+ Santa Clara Id. 14.4 8.3 6 Cumberland Bay, Isla Mas a Tierra (paratype) 13.4 7.8 5 Cumberland Bay, (holotype) 13.4 7.9 5+ Isla Mas Afuera 13.2 8.2 44 Santa Clara Id. 13.0 8.1 5+ Isla Mas Afuera 12.8 8.0 4+ Isla Mas Afuera 12.4 7.3 6 Isla Mas Afuera 11.9 7.4 5 Cumberland Bay, Isla Mas a Tierra (paratype) 11.1 6.8 44 Isla Mas Afuera 10.5 6.6 44 Cumberland Bay, Isla Mas a Tierra 10.1 6.5 54 Isla Mas Afuera 9.8 Dal. 44 Isla Mas Afuera 8.6 5.4 5+ Isla Mas Afuera 7.0 4.3 54 Cumberland Bay, Isla Mas a Tierra (paratype) 6.2 oer 5+ Cumberland Bay 5.2 3.0 44 Isla Mas Afuera Types—The holotype was collected by Dr. Waldo L. Schmitt from the littoral zone, east shore of Cumberland Bay, Isla Mas a Tierra, Juan Fernandez Islands, in 1926 (USNM_ 368900); there are 25 paratypes (USNM 679256). Synonymy— 1885 Littorina penitaria Wood, in G. Nevill, Hand List of Mollusca in the Indian Museum, Calcutta, part 2, p. 142 (San Juan Fernandez [sic]); [Nomen nudum]. 1922 Littorina mauritiana Lamarck, in N. H. Odhner, The Natural History of Juan Fernandez and Easter Island, edited by Dr. Carl Skottsberg, vol. 3, part 2, p. 223 (Isla Mas Afuera; not L. mauritiana (Lamarck, 1822). Records—ISLAS JUAN FERNANDEZ: Isla Mas Afuera (Odhner, 1922); East side of Isla Mas Afuera (Eltanin Cruise 21, Ann Cohen, Collector; USNM); Holotype and 25 para- types, east shore Cumberland Bay, Isla Mas a Tierra; North Bay, Isla Santa Clara (both collected by W. L. Schmitt in 1926; USNM). Paratypes collected by William Dodd in Del. Mus. Nat. Hist. no. 39221. [05-356 ] November 30, 1970 Littorina cincta Quoy and Gaimard, 1833 (Pls. 364, 365) Range—North, South and Stewart Islands, The Snares and Chatham Islands, New Zealand. Remarks—There is little reason for confusing the two larger species of Littorina which inhabit the shores of New Zealand. The closely brown- banded L. cincta offers a rather striking contrast Plate 364. Figs. 1-6, Littorina (Austrolittorina) cincta (Quoy and Gaimard). Figs. 1,2. Lectotype of Littorina cincta, from New Zealand (MHNP; 12.1 x 6.8 mm.). Figs. 3,4. Holotype of Littorina luctuosa Reeve (BM(NH) 1968315, 13.7 x 7.6 mm.). Figs. 5,6. Holotype of Melarhaphe zelandiae Finlay (from Transactions of the New Zealand Institute, 1926, vol. 57, pl. 18, figs. 18, 19; 17 x 10.5 mm.). Figs. 7-10. Littorina (Austrolittorina) punctata (Gmelin). Figs. 7,8. Littorina punctata, from 20 km. south of Luanda, Angola, West Africa (USNM 679288; 9.3 x 6 mm.). Figs. 9,10. Holotype of Turbo punctatus Gmelin, Senegal (from Journal de Conchyliologie, 1942, vol. 85, pl. 10, figs. 2a, 2b; MHNP, 17 x 11 mm.). Figs. 11-16. Littorina (Austrolittorina) africana (Philippi). Figs. 11,12. Lectotype of Litorina africana, from Cape of Good Hope, South Africa (Stuttgart Museum (NH): MT 106, 9.8 x 7.3 mm.; photo is from Janus, 1961, Stuttgarter Beitrage zur Naturkunde, no. 70, pl. 3, figs. 1,2). INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 473 to the lighter, grayish blue L. unifasciata anti- podum which has a single dark and often diffuse, revolving color band on each whorl. Although there is a superficial resemblance in color pattern between L. cincta and L. pintado, details of anatomy and of shell morphology confirm their distinctness. Habitat—On rocks at and above high tide line. Description—Shell reaching 20 mm. (Suter, 1913; about 0.8 inch) in length, conic-turbinate in shape, average obesity about .60 (23 specimens range from .52-.67); older individuals only moderately thick in structure, imperforate, usual- ly developing an often rather narrow flattened, crescent-shaped area adjacent to columellar cal- lus; sculptured with often faint spiral striae rein- forced by white spiral lines to a varying degree; fine to rather coarse axial lines of growth present. External color pattern consisting of the narrow, white spiral bands with wider medium to dark brown spiral bands interspersed. Aperture usual- ly dark-brown, the narrow white lines showing through inside outer lip, with a broad white band near junction of outer lip and columella. Base hardly flattened, separated from upper part of body whorl by a rather weak keel at periphery. Whorls 5-7, moderately rounded. Spire less than half the length of shell, convex, produced at an angle of about 55°. Aperture oval; outer lip rather thin, inner lip weakly concave. Columella brown to white, shallowly excavated, somewhat rimmed medially and with a flattened crescent-shaped area distally on the adjacent base. Suture im- pressed. Operculum corneus, paucispiral. Perios- tracum not evident in specimens examined. Nuclear whorls smooth and colorless; first 2-3 post-nuclear whorls dark-brown, unsculptured, Figs. 13,14. Littorina africana, from Tiger Rocks, Isipingo, Natal, South Africa (USNM 637358, 9.4 x 6.5 mm.). Figs. 15,16. Lectotype of Litorina decollata Philippi, from Natal, South Africa (Stuttgart Museum (NH): MT 107, 5.8 x 4.3 mm.; photo from Janus 1961, Stuttgarter Beitrage zur Naturkunde, No. 70, pl. 3, figs. 3,4). Figs. 17-23. Littorina (Austrolittorina) knysnaensis (Philip- pi). Figs. 17,18. Lectotype of Litorina knysnaensis from Cape of Good Hope, South Africa, near Knysna River (Stuttgart Museum (NH): MT 108, 9.3 x 6.0 mm.; photo from Stuttgarter Beitrage zur Naturkunde, no. 70, pl. 3, figs. 5,6). Figs. 19,20. Lectotype of Littorina picea Reeve [from South Africa] (BM(NH) 1968320, 6.9 x 4.5 mm.). Figs. 21-23. Littorina knysnaensis from “South Africa’; in fig. 22, basal view, and in fig. 23, note spotted color pattern particularly characteristic of base of shell in this species (USNM 633262, 11.1 x 6.9 mm.). [05-357 ] 474 Austrolittorina shining. Radula littorinid (2-1-1-1-2); similar to L. unifasciata central tooth somewhat narrow. Animal darkly pigmented on upper surfaces of tentacles, snout and foot. Verge short and thick, yellowish white in color; having a large basal flap which bears a single penial gland containing an internal hyaline accessory flagellum. Reproduc- tion unknown, probably oviparous with pelagic capsule. Measurements (mm.) length width no. whorls locality 19.2 11.0 6+ Wellington Harbour, New Zealand 18.8 10.6 6 Caroline Bay, Timaru, South Island 18.2 9.8 6+ Manukau Harbour 17.1 10.3 4+ Solander Id., Foveaux Strait 16.1 9.3 5+ Auckland 15.4 8.0 6+ Stewart Island 14.4 8.1 5+ Stewart Island 13.5 9.1 4+ Caroline Bay, Timaru, South Island 13.3 8.4 4+ Bay of Islands 11.8 7.1 5 Stewart Island es 7.1 4+ Worser Bay 9.2 6.0 4+ Stewart Island 8.8 5.4 4+ Auckland Synonymy— 1833 Littorina cincta Quoy and Gaimard, Voyage De L’Astro- labe vol. 2, part 2, p. 481, pl. 33, figs 20-21 (New Zealand); lectotype in Museum d Histoire Naturelle, Paris: 12.1 x 6.8 mm; not L. cincta Gould, 1847, Proceedings of the Boston Society of Natural History, vol, 2, p. 252 (from Puget Sound) [=L. sitchana Philippi]. 1857 Littorina luctuosa Reeve, Conchologia Iconica, Vol. 10, Littorina, pl. 13, fig. 65 (New Zealand); Holotype BM(NH) 1968315: 13.7 x 7.6 mm. 1926 Melarhaphe zelandiae Finlay, Transactions of the New Zealand Institute, vol. 57, p. 375, pl. 18, figs. 18, 19 [description and figs. = L. cincta] (Dunedin Har- bour; type in Finlay Collection, Auckland Museum, 17 x 10.5 mm.); erroneously proposed as a new species name for Littorina mauritiana ‘Lamarck’ Suter, 1913, p. 188, which = L. unifasciata anti- podum Philippi (see Finlay, 1930, Transactions of the New Zealand Institute, vol. 61, p. 224). Types—A lectotype of Littorina cincta Quoy and Gaimard is here designated from among three syntypes in the Museum National d’Histoire Na- turelle, Paris. (see pl. 364 figs. 1, 2). It is suspected that the figure in L’Astrolabe Atlas is a composite drawing as none of the syntypes match it exactly and it is smaller than the measurement accom- panying the description: 12.1 x 6.8 mm. vs 6 x 4 lines [= about 13.5 x 9.0 mm.]. The holotype of Littorina luctuosa Reeve is in the British Museum (NH) 1968315 (pl. 364, figs. 3, 4). The type of Melarhaphe zelandiae Finlay is in the Auckland Museum (pl. 364, figs. 5, 6). Joseph Rosewater Littorinidae Records—NEW ZEALAND: NORTH ISLAND: Plim- merton, Wellington (USNM); Worser Bay (USNM, MCZ); Oriental Bay, both Wellington Harbor (USNM); Wanganui (MCZ); New Plymouth (AMS); Doubtless Bay; Russell (both MCZ); Long Beach, both Bay of Islands (USNM); Hen Island, off E. coast Auckland Province (MCZ); Auckland (USNM, ANSP, MCZ); Muriwai Beach (MCZ). SOUTH ISLAND: Lyttelton; Akaroa Banks Peninsula (both AMS); Picton; Kaikoura; Timaru (all MCZ); Caroline Bay, Timaru (USNM); Kartiki, Otago (ANSP); Purakanui, N. Otago (MCZ); Hatch- ery, Portobello; Pipikariti, Dunedin; Wangaloa; Riverton (all AMS); Wangaloa; Riverton (all AMS); Point Elizabeth (MCZ); Greymouth (AMS, MCZ). FOVEAUX STRAIT: Solander Is- land (USNM). STEWART ISLAND (USNM, ANSP). CHAT- HAM ISLANDS. THE SNARES (both Suter, 1913). Littorina punctata (Gmelin, 1791) (Pls. 364, 365) Range—The Mediterranean, West and South Africa. Remarks—Although not truly a member of Indo-Pacific Littorinidae, L. punctata is a tropical species which appears to belong in the subgenus Austrolittorina and evidences many characteris- tics similar to L. unifasciata. Its shell is often subturbinate or conical, there is a tendency to Plate 365. Fig. A. Penis of Littorina cincta Quoy and Gaimard, from Plimmerton, Wellington, New Zealand (USNM 671204). Fig. B. Penis of Littorina punctata (Gmelin) from Goree, Daker, Senegal (ZMC). Fig. C. Penis of Littorina africana (Philippi) from 11 miles south of Port Shepstone, Natal, South Africa (ANSP 216678); shading indicates pigmentation noted in this and some other specimens. Fig. D. Penis of Littorina knysnaensis (Philippi) from N.E. False Bay, Cape Province, South Africa (ANSP 216643); dark area indicates pigmentation noted in some speci- mens. All drawn to same scale. [05-358] November 30, 1970 develop a semilunar depression adjacent to the columella, a white band revolving into the aper- ture, and the verge has a flap with a single hyaline spur. The range of L. punctata is quite extensive: from the Mediterranean to South Africa, via West Africa. The species may be recognized readily by the overall pattern of white spots on a dark background from which its name is derived. It is included here to avoid confusion with Indo-Pacific species whose ranges extend into or are limited to South African waters. Habitat—Shore rocks, in the spray zone. Description—Shell may exceed 18 mm. (about 0.7 inch) in length, conical to subturbinate in shape; average obesity about .64 (26 specimens range from .61-.68) moderately thick in struc- ture, imperforate, usually developing a narrow, flattened, crescent-shaped area adjacent to colu- mella callous; sculptured with only moderately well-impressed spiral striae and irregular axial growth lines. External ground color light to dark brown, with an overall pattern of white spots varying in shape from small, compact rhomboidal to elongate. Aperture medium to dark brown with a prominent white band near junction of outer lip and columella. Base somewhat flattened, sepa- rated from upper part of body whorl by a low but distinct keel at periphery. Whorls 5-7, rather straightsided. Spire less than half the length of shell, convex, produced at an angle of about 60°. Aperture oval; outer lip moderately thick, having its origin high on body whorl so that keel enters aperture; inner lip weakly concave. Columella tannish white, shallowly excavated, stout appear- am ™ a] | > he: % ae f OL. Ritts. | = | aoe ee | NEW SQhoR +, TONGA p Fel eee : “LU CALeDONia_ Os is. _ COOK] - . Norfo/k [CINCTA] 140° 160° 180° 160° Plate 366. Geographical distribution of Littorina (Austrolit- torina) cincta (Quoy and Gaimard), in New Zealand. INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 475 ing and with a narrow crescent-shaped area on adjacent base. Suture not deeply impressed. Spire sculpture often obscured by wear; 15-17 spiral striae on body whorl above keel, becoming ob- scure below on base. Operculum corneous, pau- cispiral. Periostracum not evident in specimens examined. Nuclear whorls decollate in mature specimens examined. Radula littorinid (2-1-1-1-2) similar to L. unifasciata. Animal darkly pigmented on surfaces of ten- tacles snout and foot. Verge yellowish white in color moderately short and thick with a basal flap containing a penial gland and hyaline accessory flagellum. Reproductive activity at its greatest during warm months (Tel Aviv, Israel; Palant and Fishelson, 1968), Probably oviparous and spawn- ing pelagic capsules. Measurements (mm.)— length width no. whorls locality 18.2 11.1 5+ Ghana 14.0 9.0 5+ Ghana 12.8 7.9 4+ Ghana 11.3 7.4 4+ Valencia, Spain 10.8 6.9 5+ Port Elizabeth, So. Africa 10.5 ‘Tal 4+ Valencia, Spain 9.9 6.4 5+ Alexandria, Egypt 9.0 5.8 4+ Alexandria, Egypt 8.5 5.5 4+ Alexandria, Egypt fos oye 3+ Alexandria, Egypt Synonymy— 1791 Turbo punctatus Gmelin, Systema Naturae, ed 13, vol. 1, p. 3597 (Senegalia); refers to Adanson, ‘Seneg’. Vol. 1, p. 168, t. 12, fig 1, “Le Marnat”’; 1942, E. Fischer, et al, Journal de Conchyliologie, vol. 85, p. 268, pl. 10, figs 2a, 2b; Holotype in Museum d'Histoire Na- turelle, Paris, 17 x 11 mm. E = - y | | | | | | [ | one | 4 | ee }—___|___ 4}_ 4 aaa Plate 367. Geographical distribution of Littorina (Austrolit- torina) punctata (Gmelin), in the Mediterranean, eastern At- lantic and South Africa. [05-359] 476 Austrolittorina 1826 Tricolia draparnauldii Audouin, 1826, Explication des Planches, Savigny, Description de l Egypte, Histoire Naturelle, vol. 1, pt. 4, Mollusques, p. 41, pl. 5, fig 19 (Egypt); Holotype in Museum d’Histoire Naturelle, Paris. 1845 Litorina pulchella Dunker, Zeitschrift fiir Malako- zoologie, vol. 2, p. 166 (Loanda [Luanda]); type in Berlin Museum; 1847, Philippi, Abbildungen und Beschreibungen Conchylien, vol. 2, p. 198, Litorina, pl. 4 fig 8; 1853, Dunker, Index Moll. Guineam, p. 12, pl. 2, figs 11-20. 1847 Litorina syriaca Philippi, Abbildungen und Beschrei- bungen Conchylien. vol. 2, p. 165, Litorina pl. 3, figs 21-22 (Ora Syriae [Coast of Syria]); type in Berlin Museum? 1858 Littorina guttata Reeve, Conchologia Iconica, vol. 10, Littorina, pl. 14, fig 76 (St. Vincent, Cape de Verds); not L. guttata Philippi [which is L. meleagris Potiez and Michaud]. 1932 Littorina perplexa Turton, The Marine Shells of Port Alfred, South Africa p. 133, pl. 28, fig. 960. (Port Alfred, South Africa; type in Oxford University Mu- seum). Types—Turbo punctatus Gmelin is based on Adanson’s figure of “Le Marnat’’. The specimen represented by that figure which is the holotype of T. punctatus is in the Paris Museum (pl. 364 fig. 9, 10). The holotype of T. draparnauldii Audouin is also in the Paris museum. The types of L. pulchella Dunker and L. syriaca Philippi may still be extant in the Berlin museum. Reeves L. guttata is based on Philippi’s figure of a West - lauriius AFRICANA 20° 60° Plate 368. Geographical distribution of Littorina (Austrolit- torina) africana (Philippi), in South Africa and Madagascar. Joseph Rosewater Littorinidae Indian specimen. Reeve apparently believed that it was actually the West African species, although the name L. guttata Philippi is generally held to be synonymous with the small spotted L. melea- gris. Young individuals of L. punctata appear superficially similar to L. meleagris and a com- parison of the animal characters of the two is needed. The holotype of Turton’s L. perplexa is in the Oxford University Museum. Records—SPAIN: Valencia (SMF; USNM). ITALY: Sicily (USNM). COAST OF SYRIA: (SMF). LEBANON: Beirut (USNM). ISRAEL: Tel Aviv (Palant and Fishelson, 1968). EGYPT: Alexandria (USNM). ALGERIA: Oran (USNM). MOROCCO: Melilla; Tangier; Casablanca (all USNM); Es- saouira (SMF). SENEGAL: Goree (SMF; ZMC). CAPE VERDE IDS: Mindelo, Sao Vicente (USNM). LIBERIA: Cape Palmas. FERNANDO POO: Santa Isabel. CONGO REPUB- LIC: Banana R. mouth (SMF). ANGOLA: Ambrizete (SMF); 20 km. S. of Luanda (USNM). SOUTH AFRICA: Langebaan, Saldanha Bay; Dassen Id.; Table Bay; False Bay; Hermanus; Breede R. mouth, St. Sebastian Bay; Still Bay; Mossell Bay; Knysna (all K. H. Barnard, 1963); Port Elizabeth, Algoa Bay (MCZ; USNM). Littorina africana (Philippi, 1847) (Pls. 364, 365) Range—South Africa, from the west side of Cape Peninsula, to southern Mozambique; Mada- gascar. Remarks—Littorina africana is distinct from, but apparently closely related to, L. knysnaensis. Both species occur on the south coast of Africa. The appearance of its shell and its anatomy also show similarities to other southern ocean species, L. unifasciata, L. unifasciata antipodum and fer- nandezensis and L. cincta of Australia and New Zealand. The shell of L. africana differs from these species in its development of comparatively strong spiral sculpture (although some individu- als appear almost smooth); anatomically its verge is pigmented and the basal flap relatively more pointed. Although Janus (1961) stated that the South African species L. decollata Philippi is distinct from L. africana the examination of large series and a study of their ecology would be necessary to settle questions regarding differ- ences. Material examined during the present study indicates that the two are identical. Tryon (1887) considered this to be the case and acting as first reviser chose L. africana as the senior syno- nym in spite of the obvious page priority of the name decollata. Habitat—On rocks in the splash zone. Description—Shell reaching 13.5 mm. (about .5 inch) in length, short-turbinate to subglobose in shape, average obesity about .69 (10 specimens range from .63-.77); older individuals only [05-360] November 30, 1970 moderately thick in structure, imperforate, usual- ly developing a narrow, flattened crescent-shaped area adjacent to columellar callous; sculptured with closely-spaced, raised spiral threads of vary- ing widths and irregular, sometimes coarse axial lines of growth (sometimes without strong spiral sculpture). External ground color grayish-white, with a diffuse bluish gray band encircling the body whorl and on anterior portions of spire whorls. Aperture medium to dark brown with a prominent white band near junction of outer lip and columella and another often obscure band high in aperture at junction of outer lip and body whorl. Base somewhat flattened, separated from upper part of body whorl by a low rather indis- tinct keel at periphery. Whorls 3-5, rounded; spire usually much eroded, considerably less than half the length of shell, convex, produced at an angle of about 66°. Aperture widely oval; outer lip moderately thick, having its origin quite high on body whorl, above keel, so that keel enters aper- ture. Columella usually brown with a lighter somewhat rimmed medial edge, moderately wide and excavated anteriorly, and with a flattened to excavated, brownish, crescent shaped area dis- tally on the adjacent base. Suture impressed. Sculpture consisting of ctosely-spaced, raised spiral threads of varying widths; wider threads usually separated by 2-4 narrow threads, disap- pearing on base and on eroded spire, and some- times appearing entirely smooth. Axial sculpture consisting of occasionally coarse, irregular growth lines. Operculum corneous, paucispiral. Periostracum not evident in specimens examined. Radula littorinid (2-1-1-1-2) central tooth some- what narrow similar to L. unifasciata. Animal darkly pigmented on surfaces of ten- tacles, snout, foot and on edge of mantle. Verge also darkly pigmented distally, fairly long and club-shaped; having a basal flap which bears a single penial gland containing a hyaline accesso- ry flagellum. Nothing is known concerning the reproduction and development of this species although probably it is oviparous and spawns pelagic capsule. INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 477 Measurements (mm.)— length width no. whorls locality 13:5 8.4 4 Fort Dauphin, Madagascar 10.7 7A 3+ South Africa 10.5 7.0 3+ Margate, Natal 9.9 6.7 4 Natal 9.9 tao 5 Natal 9.7 6.7 4 Natal 9.6 6.4 4 Isipingo, Natal 9.0 Bal 4 Umbhlali River, Natal 8.6 6.6 4 South Africa 8.2 5.9 3+ Inhaca Island, Mozambique re 5.0 3+ Margate, Natal Synonymy— 1847 Litorina africana Philippi, Abbildungen und Beschrei- bungen Conchylien, vol. 2, part 7, p. 199, Litorina, pl. 4, fig 10 (Caput Bonae Spei [Cape of Good Hope, South Africa]; lectotype in Stuttgart Museum (N.H.), catalogue number MT 106: 9.8 x 7.3 mm. (Janus, 1961); 1848, Krause, Die Sudafrikanischen Mollus- ken, p. 102; 1858, Reeve, Conchologia Iconica, vol. 10, Littorina, pl. 8. figs 37 a, b. 1847 Litorina decollata Philippi, ibid., p. 196, pl. 4, fig 3 (Ora Natal Africae [Coast of Natal, South Africa]); lectotype in Stuttgart Museum (N.H.), catalogue number MT 107; 5.8 x 4.3 mm. (Janus, 1961); 1848, Krause, ibid., p. 102. Types—Lectotypes of L. africana and L. decol- lata are in the Staatlichen Museum fiir Natur- kunde, Stuttgart, Germany, and were designated from Krauss material by Janus (1961). Philippi listed “Cape of Good Hope” as type locality for L. africana and “coast of Natal” for decollata. Krauss specified “Algoa Bay” for africana which may be construed as a restriction of Philippi’s type locality. Janus listed “Algoa Bay” and “Natal” as the type localities of the lectotypes of africana and decollata respectively. Records—SOUTH AFRICA: W. Cape peninsula opposite Simonstown; Robberg, Plettenberg Bay (both ANSP); Cape Recife; Port Alfred (both MCZ); East London, mouth of Nahoon River; Second Beach, Port St. Johns, Pondoland; Port Edward, Natal; Margate, 11 mi. S. Port Shepstone, Natal (all ANSP); Tiger Rocks, Isipingo, Natal (USNM; NMW); Dur- ban, Natal (MCZ; USNM): mouth of Umhlali River, Natal. MOZAMBIQUE: Inhaca Is. Delagoa Bay (both USNM). MADAGASCAR: Pointe Ibanona, Fort Dauphin (MCZ). [05-361 } 478 Austrolittorina Joseph Rosewater Littorinidae Littorina knysnaensis (Philippi, 1847) (Pls. 364, 365) Range—South Africa, from Lambert’s Bay, west coast to Natal. Remarks—The “Knysna” littorina is quite dis- tinctive and relatively easily distinguished from other South African species. “Typical” specimens exhibit a fairly prominent keel, flattened sculp- ture and brownish and tan coloration with whit- ish spots. The spots are most clearly observable above and below the wide brown peripheral color band. There is similarity, however, between the verges of L. knysnaensis and L. africana. Where differences in male anatomy are very slight, habi- tat differences or physiological barriers may oper- ate in preventing cross fertilization between the species (see Palant and Fishelson, 1968). This species occurs in part outside the range of other South African Littorina, being the dominant and/or only South African species on the west coast and becoming rare farther north on the east coast (Stephenson et al, 1940; Stephenson, 1947). It is of course similar in part to L. punctata in its spotted coloration, but differs in color pattern distribution and in shell form. The form L. africana tryphena Bartsch was based on beach worn and anomalously high- spired specimens whose appearance is very dif- ferent. Nevertheless, tryphena is an absolute syn- onym of knysnaensis. Habitat—Intertidal on rocks and in crevices of seawalls. Description—Shell reaching 13.6 mm. (about 0.5 inch) in length, turbinate in shape, average obesity about .64 (21 specimens range from .56- .70); older individuals only moderately thick in structure, imperforate, usually developing a nar- row flattened, crescent-shaped area adjacent to columellar callous; sculptured with spiral striae, between which surface of shell is flattened, and closely-spaced rather regular oblique axial lines of growth; at times spiral and axial sculpture tend to produce a reticulated pattern. External color mahogany brown in a wide band above peripher- al keel, lighter near suture and on the base, with whitish spots and streaks especially on base. Aperture dark-brown with a white band near junction of outer lip and columella. Base distinct- ly flattened, separated from upper part of body whorl by a raised keel at periphery. Whorls 3-5, somewhat flattened; spire usually eroded, less than half the length of shell, convex, produced at an angle of about 66°. Aperture oval; outer lip moderately thick, having its origin quite high on body whorl, above keel, so that keel enters aper- ture. Columella brown, but sometimes cream- colored, with a rather weakly rimmed medial edge; moderately wide and excavated anteriorly, and with a flattened to excavated, brownish to cream colored crescent shaped area distally on the adjacent base. Suture moderately impressed. Sculpture consisting of about 9 spiral striae on penultimate whorl of spire (surface of younger whorls usually to worn to count) persisting onto body whorl where 15-17 striae may be present above keel, and 9-10 below. Surface of shell between striae usually flat or only slightly raised. Axial sculpture consisting of fine, regular oblique axial lines of growth. Nuclear whorls about 3, smooth, light brown; first post nuclear whorl spirally sculptured. Operculum corneous, pauci- spiral. Periostracum not evident in specimens ex- amined. Radula littorinid (2-1-1-1-2), similar to L. unifasciata; central tooth somewhat narrow. Animal darkly pigmented on surfaces of ten- tacles, snout, and foot. Verge may also be darkly pigmented distally, moderately long and club shaped; having a pointed basal flap with a penial gland and a hyaline accessory flagellum; sperm duct open, deeply folded. Nothing is known concerning the reproduction and development of this species although probably it is oviparous and spawns pelagic capsules. Measurements (mm.)— length width no. whorls locality 13.6 7.7 4+ Algoa Bay, Cape Colony 13.0 8.0 4+ “South Africa” 12.5 8.0 4+ “South Africa” 12.2 6.9 5. Algoa Bay, Cape Colony 11.8 8.1 3+ “South Africa” 10.8 6.8 4 Muizenberg, Cape Province 10.7 6.8 44+ Cape of Good Hope 9.7 6.2 4 Margate, Natal 9.6 6.6 3+ Knysna River, Cape Province 8.7 6.0 4+ Algoa Bay, Cape Colony 8.6 5.8 5 Camps Bay, Cape Colony 8.4 5.5 3 Port Elizabeth, Cape Colony 8.0 5.3 5+ Capeland betw. Port Alfred and Cape Agulhas 7.6 4.6 4 Port Alfred, Cape Colony 6.7 4.7 Q+ Cape of Good Hope [05-362] November 30, 1970 Synonymy— 1847 Litorina knysnaensis Philippi, Abbildungen und Besch- reibungen Conchylien, vol. 2, p. 196, Litorina pl. 4, fig 4, (Caput Bonae Spei ad regionem fluminis Knysna [Cape of Good Hope, South Africa, near Knysna River]); lectotype in Stuttgart Museum (NH), catalogue number MT 108: 9.3 x 6.0 mm. (Janus, 1961); 1848, Krauss, Die Siidafrikanischen Mollus- ken, p. 102. 1857 Littorina picea Reeve, Conchologie Iconica, vol. 10, Littorina, pl. 15, fig. 83 (no locality given); lectotype BM(NH) 1968320: 6.9 x 4.5 mm. 1915 Littorina africana tryphena Bartsch, United States Na- tional Museum Bulletin 91, p. 120, pl. 38, fig. 6. (Port Alfred, South Africa); holotype USNM 187091. 1932 Littorina rietensis W. H. Turton, the Marine Shells of Port Alfred, S. Africa, p. 131, pl. 28, fig 948 (Port Alfred, South Africa); type in Oxford University Muse- um. 1932 Littorina kowiensis W. H. Turton, ibid., p. 132, pl. 28, figs. [sic] 956 (Port Alfred, South Africa); type in Oxford University Museum. 1932 Littorina africana pica ‘Reeve, W. H. Turton, ibid., p. 133 [invalid emendation of L. picea Reeve]. 1932 Littorina indistincta W. H. Turton, ibid., p. 133, pl. 28, fig 959 (Port Alfred, South Africa); type in Oxford University Museum, Types—A lectotype for Litorina knysnaensis Philippi was designated by Janus (1961) from among 8 syntypes in the Stuttgart Museum MT 108. A lectotype is here designated for Littorina picea Reeve from among 3 syntypes in the British Museum (NH): 1968320 (see pl. 364, figs. 19, 20). The type locality for L. picea is here designated as South Africa. The holotype (figured specimen) of L. africana tryphena Bartsch is in the U.S. National Museum: USNM 187091, and the para- type originally associated with the holotype has been recatalogued: USNM 664353. The types of Littorina rietensis, L. kowiensis and L. indistincta all described by Turton (1932) are in the Oxford University Museum. INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 479 Records—SOUTH AFRICA: Lambert’s Bay (Stephenson, et al, 1940); Saldanha Bay (ANSP); Dassen Id. (ZMA); Camps Bay (USNM); Chapman’s Bay, W. of Cape Peninsula (ANSP); Muizenberg (USNM); Gordons Bay, nr. Strand, False Bay (ANSP; ZMA); Simonstown (ZMC); Simons Bay, E. Cape Point (ANSP); Cape of Good Hope (USNM); N.E. of False Bay; Buffels Bay, Cape Point; Onrust, 7 mi. W. Hermanus; Hermanus; Cape Agulhas; Cape St. Blaize, Mossell Bay (all ANSP): mouth of Knysna River, Cape Province (ex. Krauss, MCZ); Beacon Isle, Plettenberg Bay (ANSP); Sea View, 16 mi. W. of Port Elizabeth (MCZ); Beacon Point, Port Elizabeth (ANSP); Algoa Bay, Port Elizabeth, Cape Colony (MCZ; USNM); Capeland, Bushman’s River mouth (USNM): Kowie R., Port Alfred (NMW); Port Alfred (ANSP; MCZ; USNM); Esplanade, East London (USNM);: Coffee Bay, 1 mi. S. of Umtata River; Port Edward, Natal (both ANSP); Second Beach, Port St. John, Pondoland; Margate, 11 mi. S. of Port Shepstone, Natal (both USNM). 20° SEYCHELLES |S. F “(§ Zanzibar 6 - 20 laurilius Reumon | is —- 40° o° Plate 369. Geographical distribution of Littorina (Austrolit- torina) knysnaensis (Philippi), in South Africa. [05-363 ] 480 Austrolittorina Joseph Rosewater [ These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence. | [05-364] Littorinidae November 30, 1970 Nodilittorina von Martens, 1897 Type: Littorina pyramidalis Quoy and Gaimard, 1833 Abbott (1954) pointed out that Nodilittorina von Martens should be considered a full genus, and designated as its tvype-species L. pyramidalis Quoy and Gaimard. He considered Nodilittorina to include also the west Atlantic species L. tuber- culata Menke with somewhat similar sculpture. Habe (1956) pointed out a difference between radulae of pyramidalis and tuberculata and on that basis proposed a new subgenus for tuber- culata, Echinolittorina, and at the same time sug- gested that Nodilittorina s.s. is an Indo-Pacific group. In the course of the present study, radulae have been studied of the various Indo-Pacific species thought to belong in Nodilittorina and all have proven close in appearance to N. pyrami- dalis. In addition to the several Indo-Pacific species belonging to Nodilittorina s.s., there are a num- ber which because of sculptural differences are here placed in the subgenus Granulilittorina Habe and Kosuge, 1966. One species, N. natalen- sis Philippi, because of its sculptural characters resembles N. tuberculata. However, its radula is not at all like Echinolittorina and, therefore, it is maintained in the genus Nodilittorina s.s. An anatomical character apparently common to members ot Nodilittorina is the partial separation of the penial gland from the basal enlargement of the penis. This condition differs from Austrolit- torina where the gland is fully incorporated in the basal enlargement. The most obvious character of Nodilittorina, the “nodose” sculpture, is quite apparent in the type-species, pyramidalis, in natalensis and nodosa. It is not always so apparent in australis with its highly variable sculpture. Nevertheless, because of similarities in anatomy and the appar- ent tendency to interbreed with nodosa it seems obvious that australis must be placedin Nodilittor- ina if current generic concepts are to be main- tained. Synonymy— 1897 Nodilittorina E. von Martens, in Weber’s Zoologische Ergebnisse Einer Reise in Niederlandisch Ost- Indien, vol. 4, pt. 1, p. 204; type-species by subse- quent designation, Abbott, 1954, p. 451: Littorina pyramidalis Quoy and Gaimard, 1833. INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 481 Subgenus Nodilittorina von Martens, 1897 Nodilittorina pyramidalis subspecies pyramidalis (Quoy and Gaimard, 1833) (Pls. 325, 326, 370, 371) Range—West coast of India and southeast Asia through the high islands of the Pacific to the Marquesas. Remarks—Nodilittorina pyramidalis is closely related to several other species of the genus in the Indo-Pacific, eastern Pacific and western Atlantic. Present means of distinguishing these species are based largely on differences in shell characters as the various species do not differ detectably in gross anatomy. When it is possible to examine such features as comparative physiology, ecology and the morphology of chromosomes additional Plate 370. Figs. 1-9. Nodilittorina pyramidalis pyramidalis Figs. 1,2. Holotype of Littorina pyramidalis Quoy and Gaimard from Jervis Bay, Australia (MHNP, 23.7 x 13.7 mm.). Figs. 3,4. Holotype of Littorina monilifera Eydoux and Souleyet, from Touranne, Cochinchina (MHNP, 11.3 x 7 mm.). Fig. 5. Lectotype of Littorina trochoides Gray (BM(NH) 87.4.26.1-8, about 10 x 7 mm.). Figs. 6,7. Nodilittorina pyramidalis pyramidalis, from Stradbroke Island, Queensland (USNM 684712, 18.5 12.8 mm.). Figs. 8,9. The same from Koh Huyong, Similan Islands, Thailand (USNM 661209, 8.6 x 5.2 mm.). Figs. 10-13. Nodilittorina pyramidalis pascua Rosewater. figs. 10,11. Holotype (USNM 679290, 13.1 x 8.8 mm.) figs. 12,13. Paratype (USNM 679291, 12.6 x 7.8 mm.) Both from Easter Island, Pacific Ocean. [05-375] 482 Nodilittorina Joseph Rosewater Littorinidae differences may be found. Arrangement of nodu- lar sculpture has been found to be quite reliable in the present study: N. pyramidalis usually displays only two rows of white nodules on the body whorl, one at the periphery and one above it with only a single row on spire whorls (the second being covered over by the succeeding whorl); N. natalensis displays three rows on the body whorl, the third row, usually slightly small- er, being located at the suture, with 2-3 rows on spire whorls and with a few raised spiral sculp- tural cords running between the rows; N. sub- nodosa shows a similar sculptural pattern, but it is much reduced at times consisting of spiral rows of weak granulations; the subspecies of N. py- ramidalis (see next species) inhabiting Easter Island is much like the nominate subspecies except that the two rows of nodules on the body whorl tend to coalesce producing axial bars rather than nodules, and this may be noted on the spire also; the west Atlantic species, N. tuberculata Menke, is like N. natalensis in possessing 3 rows of nodules on the body whorl and two on spire whorls. This definite difference between N. py- ramidalis and N. tuberculata contradicts the statement by Abbott (1954) that the two are sib- ling species as they are not strictly morphologi- cally identical. The east Pacific species N. gala- pagiensis Stearns has not been collected in suf- ficient numbers to allow adequate understanding of its variation. The latter species with three rows of nodules on the body whorl appears more closely related to the Atlantic and east African species than to N. pyramidalis. Some young in- dividuals of the east Atlantic species, miliaris Quoy and Gaimard, have only two main rows of nodules, but in later life develop multiple rows which causes me to place them with the members ~ Plate 371. Nodilittorina pyramidalis pyramidalis Quoy and Gaimard, from Koh Phi Phi, Thailand (USNM 661502). Fig. A. Radula (upper scale is 0.05 mm.). Fig. B. Penis; note partial separation of penial gland from basal enlargement (lower scale is 2 mm.). of | : L f 2mm _y of the subgenus Granulilittorina Habe and Kosuge (see world species list). Interestingly N. pyramidalis is one of those species showing an affinity for high islands or continental shores as indicated by its distribution which skirts the Pacific atolls. Perhaps its prefer- ence for high shore rocks limits the species to coasts offering such a habitat. On the coast of Queensland and New South Wales, Australia, N. pyramidalis apparently reaches its largest size (see measurements) and it was from this region (Jervis Bay) that the species was originally described. It is difficult to specify the reason for this gigantism, but it may possibly be related to a lack of optimal spawning tempera- tures or to parasitic castration either of which could prevent the onset of sexual maturity and thus prolong the active growing period. Habitat—Usually found on shore rocks con- siderably above high tide line. Description—Shell reaching 23.5 mm. (nearly | inch) in length; but usually less than 12 mm.; high cone or pyramidal in shape; average obesity about .63 (43 specimens ranging from .55 to .70); relatively thick in structure, imperforate, sculp- tured with raised spiral cords and bearing rows of raised nodules; microscopic sculpture, where not worn away or otherwise obscured, consisting of overall closely spaced wavy spiral threads. Axial sculpture consisting of oblique growth lines. Ex- ternal color, exclusive of nodules, dark reddish to blackish brown, nodules usually white, but some- times a lighter reddish brown. Aperture medium to dark reddish brown, with a narrow yellowish white band revolving inward from near anterior junction of outer lip and columella; columella also medium to dark reddish brown. Base some- what flattened; periphery nodulated. Whorls 5-6, rather flat-sided. Spire usually more than half the length of shell, produced at an angle of from 45-60°. Aperture roundly oval to nearly diamond- shaped; outer lip moderately thick; inner lip (columella) flattened, moderately excavated, es- pecially anteriorly where a broad siphonal trough is directed at an angle of 45° to anterior-posterior axis of shell. A flattened crescent shaped area on base adjacent to columella callous. Suture distinct and covering over lower row of nodules on spire whorls. Predominant sculptural feature is double row of white nodules on body whorl, each row developing 11-15 nodules; nodules often crossed by spiral cords; one row at periphery, the second just above it; nodules usually arranged one above the other, but sometimes out of phase; a single [05-376] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 483 row of nodules on spire whorls; the peripheral row on spire being covered over by next succeed- ing whorl, but sometimes just visible above su- ture. Nuclear whorls worn or decollate in all specimens examined, remaining portions smooth; postnuclear whorls nodulose. Operculum round- ly oval, paucispiral. Radula littorinoid, formula 2-1-1-1-2; central tooth narrow, tricuspid; outer marginal teeth with blunt enlargements laterally. Animal darkly pigmented on upper surfaces of tentacles, snout and foot. Verge relatively short and thickened in preserved specimens, with an enlargement near its base which is partly sepa- rated from a penial gland bearing an accessory flagellum; sperm groove deeply folded; distal end of verge appears minutely papillose. Produces a pelagic capsule having 6 spiral ridges, bearing a single egg, and measuring 160 micra in diameter (Tokioka, 1950; Habe, 1956). Measurements (mm.)— length width no. whorls locality 23.5 16.4 6 Lord Howe Id. 2h D: 14.2 6 Stradbroke Id., Queensland 21.8 14.5 6 Lord Howe Id. 19.3 Tay 6 Port Kembla, New South Wales 18.4 12.3 5 Sydney, NSW, 1k7.3 12.1 6 Stradbroke Id., Queensland 13.4 8.0 5 “Marquesas” 11.6 7.0 5+ Berhala Id., Sandakan No. Borneo 10.6 6.3 5 Pelaboean Ratoe, Java 7 5.9 5 Jamelo Bay, Luzon, P.1I. 9 6.1 5 Barrow Id., betw. Cape Dupuy and Cape Malouet, W. Australia 5.4 3.0 5 Guam, Marianas Synonymy— 1791 Trochus nodulosus Gmelin, Systema Naturae, ed. 13, vol. 1, part 6, p. 3582 (In Oceano australi, (et minor) mari; Americam meridionalem); refers to Chemnitz, Conchylien Cabinet, vol. 5, pl. 163, f. 1545, 1546; not T. nodulosus Solander, 1766 in G. Brander, Fossilia Hantoniensia, p. 10, pl. 6. 1817 Turbo trochiformis Dillwyn, A Descriptive Catalogue of Recent Shells, vol. 2, p. 826 (Southern Ocean); refers to Gmelin, T. nodulosus, p. 3582, and to Chemnitz, ibid.; not T. trochiformis Brocchi, 1814. 1833 Littorina pyramidalis Quoy and Gaimard, Voyage de L’Astrolabe, Zoologie, vol. 2, p. 482, pl. 33, figs. 12-15. (Jervis Bay [New South Wales] Australia); holotype in Paris Museum, 23.7 x 13.7 mm.; 1954, Abbott, Proceedings U.S. National Museum, vol. 103, p. 456. 1839 Littorina trochoides Gray, the Zoology of Captain Beechey’s Voyage—in His Majesty's Ship Blossom, Mollusca, p. 140 (no locality); (lectotvype, BM(NH) 87.4.26.1-8, ca. 10 x 7 mm.) 1846 Litorina vilis ‘Menke’ Philippi, Abbildungen und Besch- reibungen Conchylien, vol. 2, p. 145, Litorina, pl. 2, fig 21 (no locality). 1847 Litorina malaccana Philippi, Abbildungen und Besch- reibungen Conchylien, vol. 3, p. 15, Litorina, pl. 6, fig 17 (Pulo Pinang). 1851 Litorina cecillei Philippi, Zeitschrift fur Malakozoo- logie, 8th Jahr, p. 78 (Ryukyu Islands). 1852 Littorina monilifera Eydoux and Souleyet, Voyage sur la Bonite, Zoologie, vol. 2, p. 559, pl. 31 figs 37-39 (Touranne, Cochinchine [Viet Nam]); holotype in Paris Museum: 11.3 x 7 mm. [1950 Littorina-capsula multistriata Tokioka, Publ. Seto Marine Biological Laboratory vol. 1, no. 3, p. 151, fig 6, 2; non-binomial; is egg capsule of N. pyramidalis fide Habe, 1956, Venus, vol. 19, no. 2, p. 121.] Types—The lectotype of Trochus nodulosus Gmelin and also of T. trochiformis Dillwyn is the specimen figured by Chemnitz which may still be extant in the Zoological Museum in Copenhagen, Denmark. Holotypes of Littorina pyramidalis Quoy and Gaimard and L. monilifera Eydoux and Souleyet are in the Paris Museum. Types of species described by Philippi: L. vilis, malaccana and cecillei may be in the Berlin Museum. Nomenclature—The excellent reviews of this species by P. -H. Fischer (1967a, 1969) came to my attention when the present study was well advanced. Fischer’s decision that the oldest name for this species is N. nodulosa (Gmelin, 1791) is correct with the unavoidable qualification that Trochus nodulosus Gmelin, 1791 is preoccupied by Solander in G. Brander, 1766 (see Synonymy). The next valid name, excluding T. trochiformis Dillwyn, 1817, also preoccupied (Brocchi, 1814) is Littorina pyramidalis Quoy and Gaimard, 1833. Although one may wish to clarify nomen- clature by applying strictly the rules of priority, one cannot dispense entirely with the effects of homonymy. I agree otherwise almost entirely with Fischer’s analysis of this species, with the exception that I have not seen records from Mada- gascar or the Red Sea and Persian Gulf areas and can only suggest that Fischer's records from these localities may be based on other species, such as N. natalensis or subnodosa. Records—INDIA: Bandra, N. of Bombay (USNM): Bombay (MCZ); Vengurla, N. of Goa; Goa; Kumpta (Kumta), North Kanara (all USNM); Cape Comorin; W. of Mandapam, Gulf of Mannar; Rameswaram Island, Pamban, Palk Strait; Ramen Point, W. side of Pamban Pass, btwn. Gulf of Mannar and Palk Strait (all ANSP); Madras (MCZ, AMS, ANSP). CEYLON: W. of Kankesanturai (ANSP); Galle (RNHL, ANSP, USNM):; Columbo (MCZ, USNM); Merissa Village (ANSP); Trincomalee (YPM). THAILAND: Ko Sindarar Nua (Chance Island); Ko Huyong (South Island), Similan Islands; Laem Phan-Pha, Ko Phuket, from around Phan-Pha point; Ko Phi Phi (all USNM); Songkhla (MCZ); Ang Thong Id; Ko Tao; Ko Maprao: Sriracha (all USNM): Ko Nom Sao, Chanthaburi Province (MCZ); Ko Samet; Rayong; Lem Sing; Lem Ngob (Ngop) (all USNM). MALAYSIA: Pulau Ular, Langkawi Is- lands (USNM); Penang (MCZ); Malacca (MCZ); Pulau Anyut, Malacca Strait, just S.E. of town of Malacca (USNM); Raffles [05-377] 484 Nodilittorina Joseph Rosewater Littorinidae 80 100 120° 140 160 180° 160° ] ° | ee. ek ne or Okman 3.6 Haw, | aie bos’ ome Aan - ee Oy ace Sc ne NI Set oeas asses eee rae ere Ve FORMOSA @ | | Om p - : Hawa re ; ore wv ; ns tors é | @ sopon £ ry ae PHILIPPINE @« | é ce hy 1s. on yaa 7 MARSHALL IS. yt d § Reke oie yt Wore 5° he : are fajuro n MALDIVE » ° .? ‘ 1 Mik is m=O 2 Helen Ri | fo : Z =I BT O° GALAPAGOS ) 1s cHAGOS |” "PHOENIX 1s iS 3 TOKELAU 1S €0CO6 - KEELING ATOLLS AG PYRAMIDALIS | } NEW | 1 ZEALAND Marta Theresa ] 4. = a =—— 1 S ra n I 1 ru 80 100 yzo° 140° 160° 180 160° 140° 120" 100" BO° Plate 372. Geographical distribution of Nodilittorina pyrami- dalis pyramidalis (Quoy and Gaimard) and of its subspecies, N. pyramidalis pascua Rosewater. Light, Singapore (ANSP, USNM); Sekudu Island, Strait of Johore (USNM). VIETNAM: Ba Lang, North Annam (USNM); Ile de la Table (AMS, MCZ). CHINA: Hong Kong (ANSP, USNM); Amoy (Hsia-men); Spider Island, Foukien Province (both USNM). JAPAN: Hachijo Island, 275 mi. S. of Tokyo; Shirahama, Wakayama Pref. (both ANSP); Waki, Satsuma (BPBM). RYUKYUS: Amami Oshima (ANSP); 1 mi. N. of Shana Wan (USNM); Nago, both Okinawa (USNM, ANSP, BPBM); Zenda, Kume (MCZ, USNM, ANSP, BPBM). PHILIPPINE ISLANDS (Many localities; see map). NORTH BORNEO: Sipitang; Sulok Island, Jesselton; Berhala Island, Sandakan (all USNM); W. Marudu Bay (ANSP). INDONE- SIA: Pulau Sebesi, Sunda Strait; Dijakarta (both RNHL): Teluk Pelabuhan Ratu (MCZ, USNM); Welkomst Bay, Ban- tam (USNM); Patjitan, all Java; Bali; Larantuka, Flores Island (all RNHL). COCOS-KEELING IDS.: South Id. (Maes, 1967). AUSTRALIA: QUEENSLAND: Thursday Island, (RNHL); Murray Island, both Torres Strait; No. VI Island (both AMS): No. VIII Island, both Howick Islands (ANSP); Lizard Island (AMS); Green Island (AMS, MCZ); Fitzroy Island (AMS): Brook Island (ANSP); Palm Islands (AMS, ANSP); Bay Rock, under clumps of dead coral and rock; Cape Cleveland nr. Townsville (both USNM); Holbourne Island, off Bowen; Hayman Island, Whitsundy Passage; Lindeman Island (all AMS); Coppersmith Island, Smith Group (USNM); Brampton Island; Coquet Island (both AMS); Yepoon; N. Keppel Island, Yepoon (both AMS, MCZ); Keppel Bay; Heron Island, Capricorn Group; Bustard Bay; Noosa Heads; Caloun- dra (all AMS); Point Lookout, N.E. Stradbroke Island (ANSP, AMS); Stradbroke Island (WAM). NEW SOUTH WALES: Byron Bay; Woody Head (Wooded Bluft); Budgewoi Beach; Toukley (all AMS); The Entrance (USNM); Putty Beach (AMS); Wyargine Point, Middle Harbor, Sydney (AMS); Col- laroy (USNM); Long Reef, N. of Manly (ANSP); Port Jackson (ANSP, AMS, MCZ); Bottle and Glass Rocks, Sydney (ANSP); Coogee Beach, Sydney (USNM); btwn. Gratlo Point and Clontor, Sydney (AMS); La Perouse; Kurnell, both Botany Bay (both MCZ); Bird Island, off Wollongong (AMS); Port Kembla (USNM); Shellharbor (AMS); Jervis Bay (WAM, MHNP): Sussex Inlet Reef (AMS); Twofold Bay (MCZ, AMS). VICTORIA: Mallacoota (AMS). WESTERN AUSTRALIA: Point Gregory, N.W. end of Peron Peninsula, Shark Bay; below Quobba Light, N. of Carnarvon (both WAM); btwn, Cape Duprey and Cape Malouet, Barrow Island (WAM, USNM); mouth of False Cape Creek, La Grange Bay; 2 mi. S.W. of jetty, Broome; James Price Point, 35 mi. N. of Broome (all ANSP); Buccaneer Archipelago (AMS). NORTHERN TERRITORY: Darwin (ANSP, USNM). NEW GUINEA: Ma- nokwari; reef at S.E. entrance to Wooi Bay, Japen Island (both ANSP); Yule Island (AMS, ANSP). LORD HOWE ISLAND: (MCZ, AMS, NMW). NORFOLK ISLAND: (AMS). BONIN ISLANDS: Port Lloyd; Ani Jima (both USNM):; Mukoshima (ANSP, USNM). MARIANAS: Saipan (ANSP); Apra Bay, Guam (USNM). SAMOA: Ofu Island; Muitre Point, Tau; Fagamalo, Tau; Siulagi Point, Tau (all BPBM). TONGA: Niuafo’ou (BPBM). MARQUESAS: Hona Nui, Ua Huka; Hiva Oa; (both ANSP); Hanavave, Fatuhiva; Nukuhiva (both USNM, ANSP); Eiao (ANSP); Ua Pou; Tahuata (both USNM). Nodilittorina pyramidalis new subspecies pascua Rosewater (Pl. 370, figs. 10-13) Range—Oeno, Pitcairn, Henderson and Easter Islands, southeastern Pacific. Remarks—As mentioned in connection with the nominate subspecies, the concept of N. py- ramidalis pascua as a subspecies inhabiting the eastern-most extremities of the Indo-Pacific faunal region appears quite valid. The subspecies is distinguished primarily on the basis of its nodular sculpture. The two rows of nodules pres- ent on the whorls in pyramidalis s.s. have a tendency to coalesce in pascua forming a series of axial ridges. The presence of Nodilittorina pyramidalis on Easter Island was first mentioned by Dall (1908) and it was later recorded by both Odhner (1922) and Lamy (1936) based on separate collections. None of these workers noted the differences be- tween Easter Island populations and the species elsewhere in the Indo-Pacific. Habitat—On rocks above high tide line. [05-378 ] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 485 Description—Very similar to N. pyramidalis with exception that the double spiral row of white nodules is replaced by a row of axial bars (result- ing from coalition of juxtaposed nodules). Reach- ing 15.3 mm. in length (about .6 inch); average obesity about .64 (22 specimens range from .57 to 69). Usually only one row of small nodules on base. Axial bars on spire whorls half covered by undulating suture. Color more uniform than in pyramidalis; generally overall bluish gray, axial bars occasionally white or a dark brown; aperture very dark brown, with a dark brown flattened crescent shaped area on base adjacent to and parallel with columella. Nuclear whorls about 2, medium brown, smooth, shining; postnuclear whorls with axial sculpture. Radula and anatomy generally similar to N. pyramidalis pyramidalis. Measurements (mm.)— length width no. whorls locality 15.3 9.8 7+ All Easter Island 14.5 8.3 5+ Paratype 14.0 8.7 6+ Paratype 13.7 8.6 5+ Paratype 13.1 8.8 7 Holotype 12.8 8.2 7+ Paratype 12°3 7.8 6+ Paratype 10.1 6.2 6+ Paratype 9.3 6.3 6+ Paratype 8.9 5.8 6+ Paratype Types—The holotype of Nodilittorina pyrami- dalis pascua (a female; USNM 679290) and 24 paratypes (males and females; USNM 679291) were collected in October, 1968, on Easter Island by Pat McCoy. Additional paratypes from the same source are in the Academy of Natural Sciences, Philadelphia. The latin word pascua, meaning “Easter” is used in the combination N. pyramidalis pascua as a noun in apposition. Records—Oeno Island (USNM); Pitcairn Island (BPBM; USNM): Henderson Island (Smith, 1913); Easter Island (USNM; ANSP). Nodilittorina australis (Gray, 1826) (Pls. 325, 373) Range—Western Australia, from Esperance to Vansittart Bay. Remarks—Nodilittorina australis is endemic to Western Australia. It is closely related to another endemic species, Nodilittorina nodosa Gray, with which it is found in at least part of its range living in what appears to be the same or a very similar ecologic niche on rocks low in the intertidal zone. Comparison of gross anatomies and radulae show no outstanding differences although the shells of the two species are clearly distinct. Specimens showing characters intermediate between N. aus- tralis and nodosa found in the same population with these species are judged to be hybrids (pl. 373, figs. 11-16). The observed similarities in the anatomy of reproductive organs are believed to permit successful inter-species copulation and hybridization, although this phenomenon has not been recognized often in Mollusca (see Boss, 1964). Hybrid specimens N. australis x N. nodosa are fairly easily recognizable. They are on the average more slender than either parent (.66 obesity versus .68) tend to show more typically nodosa shell characters during early growth, but in later growth take on more the appearance of australis. True australis may be recognized by its rather globose shell, generally rough but not nodose sculpture which is often wrinkled axially, light tan to light violet colored aperture which has a relatively broad white band revolving within it. Spire sculpture is granulose but never nodulose. In comparison, the shell of nodosa is pyramidal with two rows of large whitish to orange nodules, one row at the suture and one at the periphery of the body whorl; on spire whorls there is one row at the uppermost extremity and one at the lower- most. Coloration of nodosa is dark brown, es- pecially within the aperture, with a relatively narrow revolving white band at its lower extremi- ty. The shape of aperture in australis is roundly oval, while in nodosa it is nearly diamond shaped. Related to the shape of aperture is the presence of a nodose keel in nodosa and the virtual absence of a keel in australis. Hybrid individuals exhibit a variety of characters gener- ally intermediate between australis and nodosa. Habitat—On rocks of the lower splash zone, intertidal. Description—Shell reaching 23.4 mm. (about 0.9 inch) in length, subglobose to subturbinate in shape, average obesity about .68 (28 specimens range from .63 to .75); relatively thick in struc- ture, imperforate, usually developing a flattened, crescent-shaped area adjacent to columellar cal- lous; sculptured with rather heavy spiral cords and often pronounced axial growth wrinkles. Ex- ternal color grayish to yellowish white. Aperture light yellowish tan to light violet, with a broad white band near anterior junction of outer lip and columella. Base hardly flattened with only a very low keeling effect at periphery. Whorls 4-6, mod- erately well rounded, especially the body whorl. Spire less than half the length of shell, convex, [05-379] 486 Nodilittorina Joseph Rosewater Littorinidae produced at an angle of about 75°. Aperture oval; outer lip moderately thick, inner lip concave, thickened. Columella usually light violet to tan, hardly excavated but with a flattened crescent- shaped area on the adjacent base. Anterior junc- tion of outer lip and columella projecting to form a weak siphonal channel which occasionally bears a series of very fine linear scratches. Suture not deeply impressed. Spiral sculpture varying from simple to moderately granulose, consisting of from 5-8 raised spiral cords on spire whorls and 18-20 on the body whorl; sculpture often thrown into axial wrinkles by growth pattern, either in part or over entire surface of shell— giving a reticulated appearance. Entire surface covered with closely spaced, fine, wavy, spiral threads, most noticeable in grooves between spi- ral cords; not detectable in worn specimens. Ir- regular, fine axial growth lines also present. Oper- culum corneous, only moderately thick, paucis- piral. Periostracum not evident in specimens ex- amined. Nuclear whorls about 3 in number, light tan in color, smooth; first post-nuclear whorls similarly colored but spirally sculptured, becom- ing rapidly granulose. Radula littorinid (2-1-1-1-2) central tooth very narrow (much like N. pyrami- dalis). Animal darkly pigmented on surfaces of ten- tacles, snout and foot. Verge moderately short and thick, yellowish white in color; having a bipartite (mitten-shaped) basal flap the “thumb” ap- pendage bearing a penial gland having a hyaline accessory flagellum (similar to N. pyramidalis). Nothing is known concerning reproduction and development of this species, although it probably is oviparous and spawns pelagic capsule. Plate 373. Figs. 1-4, Nodilittorina australis (Gray, 1826). Figs. 1,2. Neotype of Littorina australis Gray, from South Mole, Fremantle, Western Australia (WAM 292-70, 15.1 x 11 mm.). Figs. 3,4. An especially “rugose” specimen from Port Deni- son, Western Australia (USNM 691677, 12.3 x 8.1 mm.). Figs. 5-10. Nodilittorina nodosa (Gray, 1839). Figs. 5,6. Lectotype, from north coast of Western Australia (BM(NH) 87.4.26.10-12; 10.1 * 7.3 mm.). Figs. 7,8. Specimen from Red Bluff, Kalbarri, near mouth of Murchison River, Western Australia (USNM 691680, 13 x 8.9 mm.). Figs. 9,10. A young specimen from the same locality (7.5 x 5.2 mm.). ; Figs. 11-16. Supposed N. australia x nodosa hybrids; note early “nodose” sculpture becoming more like australis in figs. 11-14. Figs. 11,12. from Vansittart Bay, northern Western Aus- tralia (USNM 684714, 17.7 x 10.7 mm.). Figs. 13,14. from Port Denison, Western Australia (USNM 691678, 14.4 x 9.2 mm.). Figs. 15,16. Specimen from same locality as figs. 7-10, possibly a young hybrid; note elongated nodules (8.2 x 5.7 mm.) [05-380] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 487 Measurements (mm.)— length width no. whorls locality 23.4 17.6 4+ “Australia” 19.0 12.4 6 All Red Bluff, Kalbarri, West Australia 18.0 12.0 6 feo 11.0 o+ 16.2 10.2 6 16.0 abi ies 4+ 15.3 11.0 o+ 15.1 10.2 5+ 14.7 10.1 5+ 13.8 9.5 5+ 13.5 9.4 6 12.5 9.2 5+ T5 Tat 5+ 10.7 hell 5 Synonymy— 1826 Littorina australis J. E. Gray in P. P. King, Narrative of a Survey of the Intertropical and Western coasts of Australia, vol. 2, Appendix B, p. 483 (type locality here designated, South Mole [Arthur’s Head] mouth of Swan River Fremantle, Western Australia.) Ne- otype designated here: West Aust. Mus. 292-70; not Littorina australis Gray, 1839, a Risella. 1843 Litorina rugosa Menke, Molluscorum Novae Hollandiae Specimen pg. 9 (ad scopulos calcareous collis Arthurshead, od ostium fluvii cygnorum [mouth of Swan River, Fremantle, Australia]); 1844, Zeitschrift fur Malakozoologie, Jahrgang 1844, p. 57 [synon- ymizes L. rugosa with L. australis Gray, 1826]. Types—According to the notes given by Gray (1826, ibid., p. 496) specimens of L. australis were not among the species deposited by him in the British Museum collections, nor were repre- sentatives of this species found there during my recent search for types of Littorinidae. It is fairly certain, therefore, that “type-specimens of this species and also that of its only other synonym, L. rugosa Menke, may be considered lost. In order to stabilize the concept of the species I consider it necessary to designate a neotype (I.C.Z.N., Art. 10, 1961), Neotype designation—Specimens of Littorina australis Gray, 1826, were collected from rocks at South Mole [Arthur’s Head], at the mouth of the Swan River, Fremantle, Western Australia, by B. R. Wilson and A. Paterson, February 9, 1968. This is a locality from which specimens could have been collected during Captain King’s survey and it is here designated the type-locality for L. australis. It is also the type-locality of L. rugosa Menke. The specimen designated as neotype of L. australis Gray, 1826, is catalogued as Western Australian Museum 292-70 and is figured in pl. 373, figs. 1, 2. Additional specimens from the neotype lot are catalogued as U.S.N.M. 679292. Measurements of Neotype: length 15.1 mm.; width 11.0 mm. ee ee - Rep orgie - GIES oop YAva oe -KEELING DLLS Y CS Se ee Se oo° 120° 140° Plate 374. Geographical distribution of Nodilittorina australis (Gray), in Western Australia. Records—AUSTRALIA: WESTERN AUSTRALIA: Esper- ance (Hodgkin et al, 1966); Middleton Beach, nr. Albany (WAM): Foul Bay (AMS); Margaret River (WAM); Augusta (NMW); Yallingup, Cape Naturaliste (USNM, WAM); Ge- ographe Bay; Cottesloe (both AMS); Cockburn Sound (WAM); Rottnest Island (WAM, AMS): Garden Island, $.W. of Free- mantle (WAM); Freemantle (USNM); Snag Island; Beagle Islands (both WAM): Port Denison (USNM, WAM); Irwin River (AMS); Abrolhos Islands (WAM); Pelsart Islands, Geel- vink Chain, W. of Geraldton (AMS); Rat Island (WAM): Shark Bay, Freycinet Estuary, Eagle Bluff; Denham, Shark Bay, Peron Peninsula; S.E. Dirk Hartog Island (all WAM); Dirk Hartog Island, Shark Bay (AMS); below Quobba Light, Blow Holes, N. of Carnarvon (WAM); Red Bluff, Kalbarri, nr. mouth of the Murchison River (USNM, WAM); Point Cloates nr. Ningaloo (AMS); Mouth of Bigota Creek, Barrow Island; btwn. Cape Dupuy and Cape Malouet, Barrow Island, (both USNM, WAM); Vansittart Bay (USNM, AMS). Nodilittorina nodosa (Gray, 1839) (Pls. 325, 373) Range—Western Australia, from the vicinity of Geraldton northward [based on available rec- ords]. Remarks—tThe failure of this species to appear often in collections outside of Australian Muse- ums is probably due in large part to its limited distribution in Western Australia. It is also readi- ly confused with N. australis because the two species apparently hybridize and the hybrid forms look like intermediates of a variational continuum. The species is here considered to be distinct although closely related to australis. It may be recognized readily by its characteristic diamond shaped outline and the two rows of rather large whitish to reddish brown nodules on the body whorl. See remarks under N. australis. Habitat—On rocks of the splash zone, inter- tidal, usually occurs with N. australis Gray. [05-381] 488 Nodilittorina Description—Shell reaching 13.3 mm. (about 0.5 inch) in length, with a diamond-shaped out- line, average obesity about .68 (29 specimens range from .64 to .72); moderately thick in struc- ture, imperforate, usually developing a flattened crescent-shaped area adjacent to columellar cal- lous; with two rows of whitish to reddish brown nodules on spire and body whorls. External color very dark-brown between nodules. Aperture very dark-brown with a relatively narrow white band near anterior junction of outer lip and columella; white spots on inner edge of outer lip mark inner side of last nodules; columella usually lighter brown than interior of aperture. Base somewhat flattened, the effect accentuated by the large row of nodules at periphery. Whorls 4-5, rather flat- sided. Spire usually eroded, considerably less than half the length of shell, convex, produced at an angle of about 66°. Aperture diamond-shaped; outer lip moderately thick; inner lip concave, thickened. Columella usually light-brown to light-violet, very shallowly excavated, with a flat- tened crescent-shaped area distally on the ad- jacent base. Anterior junction of outer lip and columella project forming weak siphonal channel which together with inner edge of outer lip occa- sionally bear series of very fine linear scratches. Suture not deeply impressed, but undulating around nodules of preceding whorl. Spiral sculp- ture of body whorl consisting of 2 rows, each containing about 9 large nodules arranged at the suture and periphery; penultimate whorl with about 11 nodules in each of 2 rows at either suture, the anterior row partly covered over by the body whorl. Other spiral sculpture consisting of rather low cords interspersed with fine wavy white spiral threads. Axial sculpture consisting of fine irregular growth lines and crevices resulting 190° 20° \40° 160° Plate 375. Geographical distribution of Nodilittorina nodosa (Gray), in Western Australia. Joseph Rosewater Littorinidae from injury repair. Operculum corneous, rather thin, paucispiral. Periostracum not evident in specimens examined. Nuclear whorls eroded in all specimens examined. Radula littorinid (2-1-1- 1-2) central tooth extremely narrow. Anatomy of animal similar to that of N. australis. Measurements (mm.)— length width no. whorls locality 13.3 8.8 4+ All Red Bluff, Kalbarri, Western Australia 12.5 8.4 3+ 11.6 7.3 3+ 10.9 12 4 10.0 6.5 3+ 9.8 6.8 3+ 8.8 5.9 4+ 8.3 5.8 4+ 6.4 4.4 3+ 6.4 4.6 3+ Synonymy— 1839 Littorina nodosa Gray, the Zoology of Captain Beechey’s Voyage—in His Majesty’s Ship Blossom, Mollusca, p. 139 (no locality given; not figured [type locality here designated: North coast of Western Australia]); lectotype in British Museum (N. H.) 87.4.26.10-12; 10.1 x 7.3 mm. 1847, Philippi, Abbildungen und Beschreibungen Conchylien, vol. 2, Litorina, p. 160, pl. 3, fig. 7. Types—The lectotype of Littorina nodosa Gray, here designated, is in the British Museum (NH) (see pl. 373, figs. 5, 6). It bears BM(NH) catalogue number: 87.4.26.10-12; there are 2 paralectotypes under the same number. Measure- ments of the lectotype are 10.1 x 7.3 mm. Records—WESTERN AUSTRALIA: Port Denison (WAM; USNM): North Island, Abrolhos Islands (WAM); Red Bluff, Kalbarri (WAM; USNM); Quobba Point, 40 miles N. of Carnarvon (ANSP; USNM); Northwest Cape (AMS); Vansit- tart Bay (AMS; USNM). [Nodilittorina australis x N. nodosa hybrids] (Pl. 373, figs. 11-16) Range—Coextensive with N. australis and N. nodosa. Remarks—See remarks under N. australis. In shell characters hybrids appear intermediate be- tween australis and nodosa. Their average shell obesity is less than either parent: .66 compared with about .68 (23 specimens range from .61- .72). Gross anatomy of the two species is similar and hybrids show no marked differences. Labora- tory and field studies should be carried out to determine whether introgressive hybridization is taking place and other details of this phenome- non. [05-382] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 489 Habitat—On rocks of the splash zone, interti- dal. Measurements (mm.)— length width no whorls locality 18.8 11.8 5 Port Dennison 18.2 11.6 4+ All Red Bluff, Kalbarri, W. Australia 17.6 11.6 4+(eroded) 16.4 10.8 4+ 15.8 10.5 4+ 15.0 9.2 5+ 14.4 9.8 4+ 13-5 89 4+ 13.0 9.2 34+ 12.9 8.8 4+ 12.0 8.7 3+ Synonymy—So far as can be determined no scientific names have been established for these hybrid forms. The combination N. rugosa x N. nodosa has no validity or standing in zoological nomenclature. Nodilittorina natalensis (Philippi, 1847) (Pl. 376, fig. 1-6) Range—East Africa and Madagascar. Remarks—Nodilittorina natalensis has a rather narrow range in East Africa and the southwestern Indian Ocean. It is replaced in south Asia and the remainder of the Indo-Pacific by another some- what similar appearing Nodilittorina, N. pyrami- dalis. However, it may readily be distinguished by the appearance of the nodular sculpture: nata- lensis usually having at least three major spiral rows of white nodules on the body whorl and 2 or 3 rows on the spire whorls, while pyramidalis has two rows on the body whorl and one on the spire whorls; natalensis also usually has a medium- to light-brown spire apex, whereas the apex of pyra- midalis is either eroded or not noticeably differ- ent in color from the rest of the shell. In sculpture natalensis is closer in appearance to the west Atlantic N. (Echinolittorina) tuberculata than to any Indo-Pacific species. The similarity does not extend to the radula, however, that of natalensis being much better developed than that of tuber- culata whose radula appears to have suffered re- duction in the numbers of cusps on the lateral and marginal teeth and in the width of the central. Habitat—Shore rocks above the splash zone. Description—Shell reaching 14.5 mm. (about .6 inches) in length, pyramidal in shape; average obesity about .64 (32 specimens range from .59 to .69); relatively thick in structure, imperforate, 20° 40° 60° 80° 20° ns 7 MALDIVE » > is. 7g SEYCHELL:S IS. CHAGOsS |” is. 24 D Mauracs i Reumon NATALENSIS 60° 80° Plate 377. Geographical distribution of Nodilittorina natalen- sis (Philippi), in the western Indian Ocean. Plate 376. Nodilittorina (Nodilittorina) natalensis (Philippi, 1847). Note 3 rows of nodules at and above periphery. Figs. 1,2. Lectotype, from Coast of Natal, South Africa (Stuttgart Museum (NH): MT 109, 12.2 x 8.5 mm.; photo from Janus, 1961, Stuttgarter Beitrage zur Naturkunde, no. 70, pl. 3, figs. 7,8). Figs. 3,4. Specimen from Point Ibanona, Fort Dauphin, southeast Madagascar (USNM 679281, 13.1 x 7.7 mm.). Figs. 5,6. Possible paralectotypes “ex Krause’, from Natal (Senckenberg Museum, 11.5 x 7.9 mm.). [05-383] 490 Nodilittorina Joseph Rosewater sculptured with raised spiral cords and bearing rows of raised nodules; microscopic sculpture, where not worn away or obscured, consisting of overall, closely-spaced spiral threads. Axial sculpture consisting of fine lines of growth. Ex- ternal color, exclusive of nodules and apex, dark blackish brown; nodules usually grayish to yel- lowish white. Aperture medium to dark reddish brown, with a narrow white band revolving in- ward from near anterior junction of outer lip and columella, in some specimens there is a wider light colored band near posterior junction of outer lip and columella. Apex light to medium brown. Base flattened with nodulose cords. Whorls 5-7, very slightly rounded. Spire usually more than half the length of shell, occasionally nearly coe- qual with length of aperture, produced at an angle of from 53-64°. Aperture roundly oval— subquadrate; outer lip moderately thick; inner lip (columella) moderately flattened and excavated, especially anteriorly where a broad siphonal trough is directed at an angle of 45° to anterior- posterior axis of shell; a flattened, crescent- shaped area on base adjacent to columella callous. Suture rather indistinct. Predominant sculptural feature: three rows of white nodules on body whorl, each row developing 13-15 nodules often crossed by spiral cords; one row at periphery, the second just above it, the third just below suture; nodules usually arranged one above the other, but sometimes out of phase; occasionally one or two accessory nodulated rows may develop from spi- ral cords between primary rows, especially high on whorl. Two to three nodule rows appear on spire whorls. Spiral cords on base often moder- ately nodulose. In an exceedingly well-preserved specimen from Chango Island, Zanzibar (ANSP) with nuclear whorls intact, the latter consist of about two smooth, light brown volutions; first postnuclear whorl also smooth, but succeeding whorls spirally striate and becoming nodulose. Operculum thin, chitinous, light-brown, oval, paucispiral. Radula littorinid, formula 2-1-1-1-2; central tooth very narrow, tricuspid, the outer cusps held high above center cusp and close to body of tooth. Littorinidae Animal darkly pigmented on anterior upper surface of head, and also tentacles, snout and foot. Verge relatively short and thick in preserved specimens, with an enlargement near its base which is partly separated from a penial gland bearing an accessory flagellum; sperm groove deeply folded. Reproduction unknown; probably oviparous, producing pelagic capsule. Measurements (mm.)— length width no. whorls locality 14.3 9.1 6+ Fort Dauphin, Madagascar 13.7 8.4 6+ Fort Dauphin, Madagascar 13.2 7.8 7+ Fort Dauphin, Madagascar 12.2 7.3 7+ Fort Dauphin, Madagascar 11.7 7.2 6+ Coast of Natal (ex Krauss 10.5 al 6+ Coast of Natal (ex Krauss 9.6 5.9 6+ Faty, Madagascar 8.9 6.1 6+ Coast of Natal (ex Krauss 8.3 4.9 7+ Faty, Madagascar 7.4 4.9 6+ Coast of Natal (ex Krauss 6.8 4.5 4 Inhaca Island, Mozambique Synonymy— 1847 Litorina natalensis Philippi, Abbildungen und Beschrei- bungen Conchylien vol. 2, p. 160, Litorina pl. 3, fig 4. (Coast of Natal, South Africa); lectotype in Stittgart Museum, MT 109: 12.2 x 8.5 mm. (Janus, 1961); 1848 Krauss, Die Sudafrikanischen Mollusken, p. 102. Types—Janus (1961) designated a lectotype for L. natalensis Philippi from specimens collected by Krauss and deposited in the Staatlichen Muse- um fiir Naturkunde in Stuttgart (pl. 376 figs. 1, 2). Records—SOUTH AFRICA: Cape of Good Hope (ANSP); Second Beach, Port Saint Johns, Pondoland; Margate, 11 mi. S. of Port Shepstone, Natal; Port Edward, Natal; Tiger Rocks, Isipingo, Natal (all ANSP); Durban, Natal; mouth of the Umhlali River, Natal (both MCZ). MOZAMBIQUE: Inhaca Island, Delagoa Bay, (USNM, ANSP). TANZANIA: Chango (Prison) Island, W. Zanzibar (ANSP). ADEN _ PRO- TECTORATE: Conquest Bay (USNM). MADAGASCAR: Faty (Ifatz), 13 mi. N. of Tulear; Anako, 20!/2 mi. S. of Tulear; Pointe Ibanona, Port Dauphin; Flacourt, Fort Dauphin; S.W. shore Le aux Nates, S. of Ile Ste. Marie; 2.5 mi. N.E. of Pointe D’Antsiraikiraiky, N.W. Ile Ste. Marie (all MCZ). [05-384 ] November 30, 1970 Subgenus Granulilittorina Habe and Kosuge, 1966 Type: Granulilittorina millegrana (Philippi, 1848) The subgenus Granulilittorina forms a con- venient group for those rather globose Nodilit- torina having multiple rows of low granular sculpture as contrasted with the members of Nodilittorina s.s. which are more pyramidal in shape and usually exhibit only one to three rows of larger nodules. The radulae and other anatomi- cal details appear grossly similar in the two subgenera. Synonymy— 1966 Granulilittorina Habe and Kosuge, Shells of the World in Colour, vol. 2, The Tropical Pacific, p. 20, pl. 6, fig. 13; Venus, vol. 24, no. 4, pp. 313, 328; type-species by monotypy Granulilittorina philippiana Habe and Kosuge, 1966 [= Nodilittorina (Granulilittorina) mil- legrana (Philippi, 1848)]. INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 491 Nodilittorina millegrana (Philippi, 1848) (Pls. 326, 378, 379, 380) Range—From the Red Sea and western Indian Ocean islands to the western Pacific where it occurs sporadically as far east as the Marshall Islands. Remarks—Better known by several of its syno- nyms L. millegrana is an extremely variable species throughout its range in the Indo-Pacific. Much of the variation may be ecologically in- fluenced. A form exhibiting reticulate sculpture, possibly representing the doubtful species L. reticulata Anton, occurs on Indian Ocean Islands and in the Pacific has been found sporadically in the Philippines, and as far eastward as Eniwetok, Marshall islands, apparently showing a prefer- ence for islands. Axially and spirally striped color forms occur more or less randomly throughout the species range, but perhaps appear more com- monly in the Philippines, East Indies and Aus- tralia. Variation in shell granulation is consider- Plate 378. Nodilittorina (Granulilittorina) millegrana (Phi- lippi, 1848), showing shell variations. Figs. 1,2. Noumea, New Caledonia (USNM 679730, 11.8 x 7.6 mm.). Figs. 3,4. Near Jesselton, North Borneo (USNM 658369, 8.0 x 5.3 mm.). Figs. 5,10. Barrow Id., Western Australia (USNM 691690, 9.2 x 6.4 mm.). Figs. 6,7. Tsutsu, Tsushima Id., Nagasaki, Japan (USNM 679181, 8.7 x 5.5 mm.). Figs. 8,9. Point de Galle, southwest Ceylon (USNM 672392, 12.2 x 8.5 mm.). Figs. 11,12. Goa, India (USNM 442974, 10 x 6.7 mm.). Figs. 13,14. Eniwetok Id., Eniwetok Atoll, Marshall Ids. (USNM 679731, 9.3 x 6.2 mm.). [05-395] 492 Granulilittorina able. Its limits are at present considered to in- clude forms ranging from smooth to rather heavily granulose. Constant characters include the generally globose appearance, the fact that the aperture is usually more than half the length of shell and the more or less dependable presence of some degree of granulation. This is the Littorina granularis “Gray of authors” (non Gray). As shown by Yen (1942), the species granularis has as its type an Atlantic specimen and is probably synonymous with WN. miliaris (Quoy and Gaimard) from the Eastern Atlantic, rather than Tectarius muricatus (Linné) as suggested by Yen (see pl. 325). Habitat—On shore rocks above high tide line. Description—Shell reaching 13.7 mm. (about 0.5 inch) in length, subglobose to subturbinate in shape; average obesity about .67 (56 specimens range from .59 to .76); moderately thick in struc- ture, imperforate; surface occasionally smooth, but more often bearing raised, granulose spiral cords on spire and body whorls; overall micro- scopic sculpture where not worn away or other- wise obscured consisting of many fine wavy spi- ral threads most easily observable between primary granulose spiral cords. Axial sculpture usually limited to irregular axial lines of growth, occasionally, especially in island populations, granulations are in phase axially and reticulate appearance results. External color variable, rang- ing from overall grayish white to spotted and axially and spirally striped with yellow to reddish brown markings. Aperture medium to dark brown, often the former lighter color with stripes or spots of the darker color, usually with a white band revolving inward from near anterior junc- tion of outer lip and columella; columella also medium to dark brown. Base not greatly flat- tened; periphery not markedly differing in its contour from rest of body whorl. Whorls 5-7, well-rounded. Spire usually less than half the length of shell, produced at an angle of from about 68-73°. Aperture widely oval; outer lip moderately thick, occasionally slightly flaring, inner lip (columella) strongly developed, broad, thick, occasionally bulging posteriorly, shallowly excavated; with a flattened crescent shaped area on base adjacent to columella callous, at times approaching a columella chink. Suture well im- pressed. Predominant sculptural feature, 9-10 spiral cords bearing granulations on body whorl and 4-5 cords on spire whorls; granulation- bearing cords usually interspersed with non bear- ing cords, the latter often weaker; granulations Joseph Rosewater Littorinidae often subdued and some shells virtually smooth. Nuclear whorls brown to black, smooth and shin- ing; postnuclear whorls becoming rapidly granu- lose and usually having a single dark spiral color band. Operculum paucispiral, brown, oval. Radu- la littorinid, formula 2-1-1-1-2; central tooth only moderately narrow, tricuspid. Animal darkly pigmented on upper surface of tentacle, snout and foot. Verge long and slender, with an enlargement near its base partly separated from a penial gland containing an accessory fla- gellum; sperm groove deeply folded. Produces a pelagic egg capsule, having 3 tiers and an undu- late border (see Tokioka and Habe, 1953.) Measurements (mm.)— length width no. whorls locality 13.7 9.2 5 Goa, India 13.7 8.1 5 Bombay, India 12.3 8.6 4+ Pt. de Galle, Ceylon 11.3 7.3 5 Pondicherry, India 10.7 7.2 4 Goa, India 9.7 6.3 5 Koh Phi Phi, Thailand 8.4 5.4 6 Souillac, Mauritius 8.1 5.6 5 Hong Kong 7.8 5.4 4 Barrow Island, Western Australia 6.7 4.2 4 Réunion 5.9 4.5 5 Keppel Bay, Queensland 5.5 3.6 4 Koh Huyong, Thailand Synonymy— —Littorina granularis ‘Gray’ of Authors, non Gray, 1839 [Gray’s holotype, BM(NH) 87.4.26.9 is an eastern Atlantic species, Nodilittorina miliaris (Quoy and Gaimard]. 1847 Litorina picta marmorata Philippi, Abbildungen und Beschreibungen Conchylien, vol. 2, Litorina, p. 167, pl. 3, fig. 26 [lectotype figure] (provincia Ilocos borealis insulae Luzon); not L. marmorata Pfeiffer, 1839. 1847 (Sept.) Litorina ventricosa Philippi, ibid., vol. 3, Litorina p. 51, pl. 6, fig 19 [lectotype figure] (Pulo Pinang [Penang Island, Malaysia]); not L. scabra ventricosa Philippi, 1847 (April). 1848 Litorina millegrana Philippi, ibid., p. 65, pl. 7, fig. 15 [lectotype figure] (Red Sea), original measurements about 13 x 11 mm. 1852 Littorina radiata Eydoux and Souleyet, Voyage sur la Bonite, vol. 2, p. 562, pl. 31 figs. 46, 47 (Touranne, Cochinchine [Viet Nam]); lectotype BM(NH) 54. 7.24.389, 11.5 x 7.6 mm. 1857 Littorina novaezelandiae Reeve, Conchologia Iconica, vol. 10, Littorina pl. 14, fig. 74 (New Zealand [Trin- comali, Ceylon, here corrected]). lectotype BM(NH) 1966124, 13.2 x 9.0 mm. designated by Biggs, 1966, Journal of Conchology, vol. 26, no. 2, p. 138, pl. 7, fig 5. 1857 Littorina granocostata Reeve, ibid., pl. 15, fig 79 (Bris- bane Water [New South Wales] Australia); lectotype BM(NH) 1968318, 6.3 x 4.4 mm. 21859 Littorina vidua Gould, Proceedings of the Boston Society of Natural History, vol. 7, p. 138. (Ousima [= Amami-o-shima, Ryukyu Islands, according to John- son, 1964]); type lost. [05-396] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 493 i Plate 379. Nodilittorina (Granulilittorina) millegrana (Phi- lippi, 1848). Fig. A. Radula of specimen from Aden (USNM 679341; lower scale is 0.05 mm.). Fig. B. Penis of specimen from Sekudu Id, near Singapore, Malaysia (USNM 660756; upper scale is 1 mm.). 1876 Littorina melanacme E. A. Smith, Journal of the Linnean Society of London, Zoology, vol. 12, p. 552, pl. 30, fig. 21 (San Christoval, Solomon Islands); Holotype BM(NH) 76.1.10.67, 9.6 x 6.1 mm. 1885 Littorina erronea Nevill, Hand List of Mollusca in the Indian Museum, part 2, p. 152 (Balapiti, Ceylon); new name for Littorina novaezealandiae Reeve, 1857. 1887 Littorina granicostata E. A. Smith, Proceedings of the Zoological Society of London for 1887, part 3, p. 519, fig. 2, (Christmas Island, Indian Ocean); Holotype BM(NH) 1968358, 10.4 x 7.2 mm.; is Littorina insu- laris E. A. Smith, 1889. 1889 Littorina insularis E. A. Smith, Proceedings of the Zoological Society of London for 1888, part 4, p. 536; new name for L. granicostata E. A. Smith, 1887. 1897 Littorina ventricosa strubelli von Martens, Zoologische Ergebnisse Einer Reise in Niederlandisch Ost- Indien, vol. 4, part 1, p. 208 (Krakatau, Indonesia); type in Zoological Museum Amsterdam ?; refers to L. pusilla, and to Philippi, “Abbild. Neuer Conch. vol. 2, p. 164, pl. 4, fig 15 [pl. 3, fig 23?] and to Kuster, Conchylien-Cabinet, vol. 2, pt. 9, p. 11, p. 1 fig. 20-22; 1925 Jutting, Treubia Batavia, vol. 6, p. 142 [not a new variety of Jutting’s as indicated by H. B. Preston, Zoological Record, vol. 62, Mollusca, p. 54]. 1915 Littorina eudeli Sowerby; Annals and Magazine of Natural History, series 8, vol. 16, No. 93, p. 167, pl. 10, fig 5 (Pondicherry, India); helotype BM(NH) 1919, 12.31.33,12x 74mm. Lch; 1936 Littorina chaoi T-c Yen, Notes de Malacologie chinoise, vol. 1, fas. 3, Musee Heude, Shanghai, p. 3 (Pok-hoy {Gulf of Tonkin] China); 1937, ibid., fas. 4, figs 2, 2a, 2b; holotype in Museum Heude? Plate 380. Type specimens of synonyms of Nodilittorina (Granulilittorina) millegrana Philippi, 1848). Fig. 1. Lectotype figure of Litorina millegrana Philippi, from the Red Sea (from Abbildungen and Beschreibungen Conchylien, vol. 3, Litorina, pl. 7, fig. 15, about 13 x 11 min.). Figs. 2,3. Lectotype of Littorina radiata Eydoux and Soul- eyet, from Touranne, Viet Nam (BM(NH) 54.7.24.389; Lib X76 mm:): Figs. 4,5. Lectotype of Littorina novaezelandiae Reeve, a smooth specimen from Trincomali, Ceylon (BM(NH) 1966124; 13.2 x 9 mm.). Figs. 6,7. Holotype of Littorina melanacme E. A. Smith, from San Cristobal, Solomon Ids. (BM(NH) 76.1.10.67; 9.6 x 6.1 mm.). Figs. 8,9. Holotype of Littorina insularis E. A. Smith [is also holotype of L. granicostata Smith] from Christmas Id; Indian Ocean (BM(NH) 1968358, 10.4 x 7.2 mm.). Fig. 10. Holotype of Littorina urieli Biggs, from Eilat, Israel (BM(NH) 1966121, 7.3 x 4.5 mm.). Figs. 11,12. Lectotype of Littorina granocostata Reeve, from Brisbane Water, Australia (BM(NH) 1968318, 6.3 x 4.4 mm.). lee Fes Figs. 13,14. Holotype of Littorina eudeli Sowerby, from Pondicherry, India (BM(NH) 1919. 12.31.33, 12 x 7.4 mm.). 1953 [Littorina-capsula hagruma Tokioka and Habe, Publica- tions of the Seto Marine Biological Laboratory, vol. 3, no. 1, pp. 55, 56 (Tanabe Bay, Japan); Habe, 1956, Venus, vol. 19, no. 2, pp. 117-121, fig. B; non- binomial]. 1966 Granulilittorina philippiana Habe and Kosuge, Venus, vol. 24, no. 4, pp. 313, 328, [figured in] Habe and Kosuge, 1966, Shells of the World in Colour, Hoikusha, vol. 2, p. 20, pl. 6, fig 13 (Goza, Shima Peninsula, Honshu, Japan); Holotype in National Science Museum, Tokyo, 6.8 x 5.2 mm. 1966 Littorina urieli Biggs, Journal of Conchology, vol. 26, p. 137, pl. 7, figs 1, 2 (Bilat, Gulf of Eilat, northern end Gulf of Aqaba, Israel) Holotype, BM(NH) 1966121, 7.4 X 5.0 mm. Types—The types of species described by Phi- lippi in Abbildungen und Beschreibungen Con- chylien should be in the Berlin Museum, al- though I have not been able to confirm their presence. Until this is possible Philippi’s figures [05-397] 494 Granulilittorina Joseph Rosewater Littorinidae must be considered accurate representations of the species, and are here held to be the lectotypes. It is interesting to note that the lectotype (figured specimen) of L. radiata Eydoux and Souleyet is in the British Museum (N.H.), while only two paralectotypes were found at the Paris Museum. Although it would be expected that the ‘Bonite’ collections would be in the Paris Muse- um exclusively, a large number of the mollusk types of Eydoux and Souleyet were bequeathed by Souleyet to the British Museum (NH) (see Gray, 1855). A lectotype for Littorina novaezelandiae Reeve was designated by Biggs (1966) and the type locality is here corrected from New Zealand to Trincomali, Ceylon. Other types of Reeve, Smith, Sowerby and Biggs are in the British Museum (NH). The type of Littorina chaoi Yen may be in the Museum Heude, Shanghai. The holotype of Gran- ulilittorina philippiana Habe and Kosuge is in the National Science Museum, Tokyo. Nomenclature—The problem of selecting a proper name for this species from among possible synonyms is compounded because of its variabil- ity, the fact that somewhat similar species occur elsewhere in the oceans, and because some early workers neglected to cite type localities with their species descriptions. The earliest valid name ap- Plate 381. Figs. 1,2. Holotype of Littorina miliaris Quoy and Gaimard, 1833, from Ascension Id., Atlantic Ocean (MHNP, 14.6 x 9.4 mm.). Figs. 3,4. Holotype of Littorina granularis Gray, 1839 [east- ern Atlantic] (BM(NH) 87.4.26.9, 14.2 x 10 mm.). pears to be L. millegrana Philippi, 1848. (see synonymy). The tamiliar name L. granularis Gray, 1839, has as its holotype a worn specimen which was interpreted by Yen (1942) as being Tectarius muricatus Linné. I believe it to be actually a worn specimen of Nodilittorina miliar- is (Quoy and Gaimard) a west African species. There is also a possibility that L. reticulata Anton, 1839, may figure in the competition for the oldest name for this species. However, reticulata like granularis Gray, has no type locality and the only illustration, that given by Philippi and said to be of Anton’s type shows a specimen which is impossible to identify with certainty. Littorina picta marmorata, usually considered to be a vari- ant of the Hawaiian species picta Philippi, but actually described from the Philippines, and L. ventricosa Philippi, are both junior homonyms and therefore fail to qualify as valid names, leaving millegrana as the next available taxon. Records—RED SEA: Eilat, Israel (G. Frankel Coll.). GULF OF ADEN: Aden (ZMC). MADAGASCAR: S.W. Shore Ile des Nattes, S. of Ile Ste. Marie; Ste. Luce, S.E. Madagascar (both USNM). INDIAN OCEAN ISLANDS: E.. of Souillac (Savanne R.), Mauritius; Caves Point, W. Mauritius; Réunion; Dunidu Id., N. Male Id., Maldives; Fadiffolu Atoll, (all USNM). INDIA: Bandra, N. of Bombay; Bombay; Goa; Cape Comorin (all USNM); Pondicherry (USNM, AMS). CEYLON: S. Shore Fort Frederick, Trincomalee; Pt. de Galle (both USNM). THAILAND: Laem Phan-Pha, Koh Phuket; Koh Huyong, Similan Ids.; Koh Phi Phi (all USNM):; Songkla (MCZ); Koh Maprao; Koh Tao (both USNM); Prachaup Khiri Khan (MCZ); Koh Nom Sao, Chantaburi Province (MCZ); Koh Sichang; Koh Chala; Rayong; Koh Kut (all USNM). MALAYSIA: Penang (MCZ); Batu Ferringgi, Penang (MCZ); Pulau Ular, Langkawi; Pulau Jerak, W. of Sembilan Ids.; Pulau Anyut, Malacca Str.; Pulau Besar, Malacca Str. (all USNM); Raffles Light, Singapore (USNM, ANSP); Sekudu Id., Johore Str. (USNM); CHINA: Hong Kong; Big Wave Bay, Hong Kong. VIETNAM: (USNM). RYUKYUS: Odomari, Oki- nawa, (USNM). JAPAN: Tsutsu, Tsushima Id., Nagasaki Pref., Kyushu (USNM); Miura Peninsula and Goza, Shima Peninsula, Honshu (both Habe and Kosuge, 1966). PHILIP- PINES: Santo Domingo, Batan; Jamelo Bay; S. Shore, Fort Mills, Cavite; Maricaban Id. (all USNM); Corregidor Id. (USNM; ANSP): Mariveles; Grand Is., Subic Bay; Bolinao Bay; Lingayen, all Luzon (all USNM); Marinduque; Pt. Naso, Panay; Zamboanga, Mindanao; Balabac; Bucas Id.; Jolo; Tumindao (all USNM). INDONESIA: Keledjitan, Bantam, Java; Pelabuhan Ratu, Preager, Java; Krakatoa (ZMC); Sulok Id., Jesselton, N. Borneo; Morotai, Halmahera Group, East Indies, Moluccas (all USNM). COCOS-KEELING ATOLLS: (ANSP). AUSTRALIA: QUEENSLAND: Lizard Id. (AMS); Los Isles (USNM); Halfmoon Bay, nr. Cairns; Fitzroy Id.; Palm Id. (all AMS); Bay Rock (USNM); Royal Seaforth, nr. Lindeman Id.; Brampton Id.; Heron Id., Capricorn Grp. (all AMS); Keppel Bay (USNM); North Keppel Id.; Barney Pt., Port Curtis; Lady Elliot Id.; Bundaberg; Point Vernon, Her- vey Bay; Caloundra (all AMS); Cape Cleveland (USNM). NEW SOUTH WALES: nr. Wollongong. WESTERN AUS- TRALIA: btwn. Cape Dupuy and Cape Malouet, Barrow Id.; mouth Bigota Creek, W. side Barrow Id.; Airport Beach, Barrow Id.; Broome (all USNM). MELANESIA: S.E. En- trance Wooi Bay, Japen Id., West Irian (ANSP); Karkar Id., North-East New Guinea; Matupi Isl. Rabaul, Bismarks; Makira Hbr., San Cristobal, Solomons; Taden Reef, N. of Heinghene, New Caledonia (all AMS); N. of Touho, New Caledonia; E. of Nani Id. (both USNM); Noumea, New Caledonia (AMS, USNM); Suva; Irvines, nr. Malaqereqere, both VitiLevu, Fiji (both USNM). PALAU IDS: Melekeiok, Babelthaup Id. (USNM). MARSHALL IDS.: Eniwetok (USNM). [05-398 ] November 30, 1970 —— —~-B ! 3 | 38 9 i} fu ' Pe = ' t3_ ee =e 3 3 8 \ i) t f 3s f ie 8 = ' iy 4 ‘ a \ ' ' ' ' & 1 = = = BS \ ' i} a ie 8 q 1s w \ x \ 3 a 8 @ x ' a Zw resi $ \ $ % woe - [ee | i ! *e, ° a? BA ' < > 37 2 ° { & = to °. ° 1 2 S4,%, zu ° ——— x eae Ams zi —{8 3! r og 2. ¥ fa et : s | g 1 Pan \« i i |z ' < 1 wh a oa G Zo} z te Zatl 38 7 O25 ports 8 \ Ss oO ! ' < ' ‘ 5 9 ey ! \ af fe TRS t +_—__ er coe 4? bal YS i) } i = H \ vA 7 w ' ; Ty 1 z \ i) a i) SF 2) a8 ! 4S i a \ aa a z er ae ' ghia oe, 1 IS Felt @ i a i ° Cb 8 ! % | = < — ig Shy i { oo ! 4, \ as ' | Mi Hera ! ae 1 i} NED H ' ' \ H ' ‘| | , 8 == , + 3 } ¢ 3 ' \ e 4 ' + seu eu ' z © to | 4 f Ie O82 8X ' | a 23 < 3 { } « mo OG r4 1 2 \ ° S st = -9 i \ fi: a | m\y t ay H s Bh q 2 H A ee S$ \ fy ! 7 \ | : | e (eee : | : ie FAdhal \ z 8 © 1 = § = ! lo Pa s| | __g | em , wi ° Maes peas re i I 3 7 S ite <= iN’ t ' _y i as Rea ' z 1 } 1 i} ! 1 4 H H ° | Fs co} s 8 © ° s 2. S Plate 382. Geographical distribution of Nodilittorina (Granu- lilittorina) millegrana (Philippi), in the Indian Ocean and western Pacific, and of N. (G.) picta and N. (G.) cinerea (Pease), in Hawaii and the Marquesas Islands respectively. INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 495 Nodilittorina subnodosa (Philippi, 1847) (Pl. 383, figs. 1-10) Range—Red Sea and Persian Gulf areas. Remarks—Nodilittorina subnodosa is an ex- ceedingly variable species within its rather nar- row area of habitation in the Red Sea and Persian Gulf. The variation is expressed particularly in the degree of shell granulation, different in- dividuals ranging from distinctly nodulose to hardly sculptured. One character apparently rare in Nodilittorina, but present in this species, is the white columella, which serves to distinguish subnodosa from natalensis, its nearest geographic relative. The relationships of subnodosa are somewhat obscure. Certain individuals from Per- sian Gulf populations resemble in part the east- ern Atlantic N. miliaris Quoy and Gaimard (Rev. H. E. Biggs, personal communication, 1968). I consider that the resemblance is superficial, and that the two are quite distinct. I believe, however, that subnodosa may possibly share a common ancestry with N. miliaris. The two Persian Gulf populations examined indicate that N. sub- nodosa is more weakly sculptured and may reach a larger size there than in the Red Sea. However, considerable additional comparative material is needed to prove this trend in shell morphology. Habitat—Shore rocks above high tide line. Description—Shell reaching 13.8 mm. (about My inch) in length, pyramidal to pyramidal- turbinate in shape; average obesity about .66 (38 specimens range from .60 - .70); relatively thick in structure, usually imperforate with some larger and older individuals developing small umbilical opening; sculpture varying from weakly granu- lose to only moderately nodulose; microscopic sculpture where not worn away consisting of fine closely spaced, wavy spiral threads; axial sculp- ture consisting of fine, irregular lines of growth. External color, exclusive of nodules and apex, yellowish to pinkish white, occasionally spotted dark brown between nodules; nodules white. Aperture reddish orange to dark brown, with a narrow white band revolving inward from near anterior junction of outer lip and columella; white band obscure in some specimens. Dark apertural coloration usually ending at anterior junction of outer lip and columella; columella white. Apex light to medium brown. Base moder- ately to considerably flattened, sculptured with spiral, granulose cords. Whorls 5-7, hardly to moderately rounded. Spire usually more than half the length of shell, produced at an angle of from [05-399] 496 Granulilittorina 55-62°. Aperture oval, outer lip moderately thick, tending to be slightly shouldered at the suture; inner lip (columella) thick and moderately flat- tened; shallowly excavated anteriorly; a flattened, crescent shaped area on base adjacent to columel- la and an umbilical opening occasionally present in older individuals. Suture indistinct only in more nodulose specimens. Predominant sculp- ture: three spiral rows of medium sized nodules above periphery on body whorl, often with addi- tional rows of smaller nodules between principal rows; usually 1-3 principal rows of nodules on spire whorls with rows of smaller granulations between. Sculpture in some specimens severely muted, consisting only of spiral rows of granula- tions showing little or no size separation. Nuclear whorls worn or decollate in all specimens ex- amined, light brown in color; postnuclear whorls spirally nodulose. Operculum thin, chitinous, light brown, paucispiral. Anatomy grossly similar to N. natalensis. Re- production unknown. Records—RED SEA: (MCZ); N.E. Museri Id., Dahlak Archipelago, Ethiopia (RNHL); Eilat, Gulf of Aqaba (USNM). PERSIAN GULF: Bushire, Iran (ZMC); near Abu Dhabi, Trucial Coast (USNM). Joseph Rosewater Littorinidae Types—tThe location of the type specimens of Litorina subnodosa Philippi is not definitely known, although it is probable that they are in the Berlin Museum. Of the two specimens repre- sented in Philippi’s illustration, the larger, his fig. 9, is here designated as the lectotype (see pl. 383, fe. 2) Measurements (mm.)— length width no. whorls locality 13.8 8.8 5+ Abu Dhabi, Trucial Coast 12.8 8.3 5+ Abu Dhabi, Trucial Coast 12.3 8.6 5+ Abu Dhabi, Trucial Coast 11.9 8.0 6 Abu Dhabi, Trucial Coast 1.1 7.2 5+ Abu Dhabi, Trucial Coast 10.4 7.0 4+ Abu Dhabi, Trucial Coast 9.7 6.8 4+ Abu Dhabi, Trucial Coast 9.0 6.3 4+ Abu Dhabi, Trucial Coast 8.4 5.3 7 Eilat, Gulf of Aqaba tee 4.3 6 Eilat, Gulf of Aqaba 6.2 4.2 6 Eilat, Gulf of Aqaba 5.2 3.5 5+ Eilat, Gulf of Aqaba Synonymy— 1847 Litorina subnodosa Philippi, Abbildungen und Besch- reibungen Conchylien, vol. 2, p. 161, Litorina, pl. 3, figs 8, 9 (Red Sea). Plate 383. Nodilittorina (Granulilittorina) subnodosa. (Phi- lippi, 1847) Fig. 1. Lectotype figure of Litorina subnodosa Philippi, from the Red Sea (from Abbildungen and Beschreibungen Conchylien, vol. 2, Litorina, pl. 3, fig. 9, about 19 x 12 mm.). Figs. 2,3. Specimens from an old collection from Gulf ot Suez (USNM 23233, 14.5 x 9.6 mm.). Fig. 4. Rather smooth specimen from near Abu Dhabi, Trucial Oman Coast, Persian Gulf (USNM 669124, 11.8 x 7.8 mm.). Figs. 5,6. Bushire, Iran, eastern Persian Gulf (USNM 679285, 8.2 x 5.3 mm.). Figs. 7-10. Strongly nodulose to only moderately granulose specimens from Eilat, Gulf of Aqaba, Red Sea (USNM 671239, 7,9: 8.3 x 5.2 mm.; 8,10: 7.5 x 4.9 mm.). [05-400] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 497 20° 40° 60° 80° eos ae ae tec — — 20° MALDIVE » © 1s. - | SEYCHELLES IS. ‘ 1 | Plate 383a. Geographical distribution of Nodilittorina (Granu- lilittorina) subnodosa (Philippi) in the Red Sea and Persian Gulf. Nodilittorina leucosticta subspecies leucosticta (Philippi, 1847) (Pl. 384, figs. 1-5) Range—India and Ceylon. Remarks—It is extremely difficult to quantify the differences between G. leucosticta and G. millegrana and on occasion it has seemed impos- sible to maintain them as separate species. Never- theless populations of leucosticta occur together with millegrana in which the individuals be- longing to each species are clearly separable. Outside the supposed range of leucosticta, no Plate 384. Nodilittorina (Granulilittorina) leucosticta and sub- species. Figs. 1-5. N. leucosticta leucosticta (Philippi). Fig. 1. Lectotype figure of Litorina leucosticta Philippi [Bombay, India] (from Abbildungen and Beschreibungen Conchylien, vol. 2, Litorina pl. 3, fig. 11, about 8 x 6 mm.). Figs. 2,3. Pointe de Galle, southwestern Ceylon (USNM 672389, 9.5 x 6.2 mm.). Figs. 4,5. Goa, India (USNM 442970, 13.1 x 7.8 mm.). Figs. 6-13. N. leucosticta biangulata (von Martens). ~ Figs. 6,7. Holotype of Littorina biangulata von Martens, from Benkulen [southwest] Sumatra (ZMA, 11 x 7.1 mm.). Figs. 8,9. Pulau Nias, Mentawai Islands. southwest of Sumatra (USNM 654444, 6.6 x 4.1 mm.). Figs. 10,11. Mouth Hienghene River, Hienghene Area, New Caledonia (USNM 637368, 10 x 6.3 mm.). Figs. 12,13. West tip Corregidor Id, Luzon, Philippines (USNM 637360, 9 x 5.5 mm.). Figs. 14-19. N. leucosticta feejeensis (Reeve). Figs. 14,15. Lectotype of Littorina feejeensis Reeve, (BM(NH) 1968319, 7.1 x 5 mm.). Figs. 16,17. Ovatoa, northwestern Vanua Levu, Fiji (USNM 694772, 7.1 x 4.7 mm.). Figs. 18,19. Tutuila, Samoa (USNM 488720, 7.2 x 4.7 mm.). [05-401] 498 Granulilittorina such mixing of phenotypes occurs. It is, therefore, fairly apparent that G. leucosticta is a valid species with more restricted range than mill- egrana and that the two are clearly separable on shell morphology alone. The shell of leucosticta averages more slender than millegrana (obesity .65 vs. .67). The single most. striking feature in Jleucosticta is the shouldered appearance of the body whorl in mature individuals. The pattern of hardly raised white nodules on the spiral cords is also distinc- tive as these are usually interrupted with brown spots; in some populations the shell ground color is dark gray to brown, and here the white semi- nodulated markings are very prominent. The markings are always less pronounced, however, than in N. pyramidalis. The aperture in leucostic- ta usually is proportionately more than half the length of shell except in large and apparently anomalous individuals which are not rare in collections. The subspecies leucosticta biangulata von Martens and lI. feejeensis Reeve replace leucostic- ta s.s. in the East Indies and western Melanesia and in the Pacific Islands respectively. Differ- ences between the subspecies are of a consider- ably qualitative nature and involve a shift in emphasis of intensity of expressed sculpture and coloration. If these taxa are to be considered subspecies rather than distinct species or even portions of a variational continuum, one must refer to specimens from the several geographical areas cited and to the descriptions of the various morphological peculiarities of each. In such cases species and subspecies concepts are considerably strained. Habitat—Intertidal on rocks. Description—Shell reaching 16.2 mm. (about °/s inch) in length, turbinate to elongate oval in shape; average obesity about .65 (22 specimens range from .59 to .70); only moderately thick in structure; imperforate; surface sculptured with rather low and closely spaced spiral cords 2-4 of which, usually three, on body whorl, are stronger than the rest; cords bearing low white, often elongate nodules separated by brown interspaces; shell shouldered about #/3 length of body whorl from suture to first strong cord; details of color and sculpture becoming obscured in larger in- dividuals; overall microscopic sculpture where not worn away or otherwise obscured consisting of fine wavy spiral threads. Axial sculpture lim- ited to irregular, often closely spaced lines of growth. External color variable, ranging from Joseph Rosewater Littorinidae grayish white base color to a dark gray or brown, usually with the low white nodules offering dis- tinct contrast. Aperture medium to dark brown, with a white band revolving inward near anterior junction of outer lip and columella, with occa- sional fainter bands posteriorly; columella usual- ly lighter colored than rest of aperture. Base slightly flattened; periphery often marked by po- sition of one of stronger spiral cords. Whorls 5-6; spire whorls only moderately rounded; body whorl rather flatsided at its center and shoul- dered. Spire usually less than half the length of shell, produced at an angle of from about 62-70°. Aperture elongate oval, outer lip only moderately thick; inner lip strongly produced, rather straight, only occasionally slightly bulging posteriorly, with a flattened crescent shaped area on base adjacent to columella callous, Suture well impressed. Body whorl subtly shouldered; pre- dominant sculptural feature: about three well spaced, low spiral cords on body whorl; cords with low white, sometimes elongate nodules, separated by brown color spots; sculpture char- acteristics often worn away or not evident in older individuals. When present, nuclear whorls about 1*/2-2, light brown, smooth; postnuclear whorls becoming darker brown and spirally sculptured. Operculum paucispiral, brown, oval. Radula lit- torinoid, central tooth only moderately narrow tricuspid. Animal darkly pigmented on upper surfaces of tentacles, snout and foot. Verge of the Nodilit- torina type, with a basal enlargement and _par- tially separated penial gland with an accessory flagellum; main portion of verge relatively long and slender, with an open deeply folded sperm duct along its posterior-medial edge. Nothing is known concerning the reproduction and develop- ment of this species, although it is suspected to involve a pelagic capsule as yet undescribed. Measurements (mm.)— length width no. whorls locality 16.2 9.8 5+ All Bombay, India 15.1 9.3 5+ 14.7 8.7 4+ 13S 8.3 4+ 11.0 7.1 3+ 10.7 7.2 4+ 9.8 6.8 4+ 8.7 6.1 3+ 7.9 5.2 4+ 6.6 4.5 3+ Synonymy— 1847 Litorina leucosticta Philippi, Abbildungen und Besch- reibungen Conchylien, vol. 2, Litorina, p. 162, pl. 3, [05-402] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 499 80° 100° 120° 140° 160° 180° 160° lac SO T Sa ——T T — - 4 5)? Hecho | Bie ) sd | lala VF ownane tise ONIN ESTs ae IG amaan- arias | arena eae tar? @\ NN ign ne q 3 1s 2 \d iy L,| PHILIBPINE ov a mA | BV L oo | Ulthe Eniwetok 8 MARSHALL IS. WLI oom | spe | oY | of ) aed Mejuro | 2 mie OS Ne carouine ~e Palmyra fo % es or A as ie fi = 4 ss = rea oa |S Chrisimes ig SFO crane’ Sy Te % ‘ 2 ge orien ts . = ae “ouinensy LF % oy, @ ‘PHOENIX 15 <3 7 at “en Javea TOKELAU IS 1 Plate 385. Geographical distribution of Nodilittorina (Granu- lilittorina) leucosticta leucosticta (Philippi) and of its other two subspecies, N. (G.) leucosticta biangulata (von Martens) and N. (G.) leucosticta feejeensis (Reeve). fig 11 [lectotype figure; original measurement about 8 x 6 mm.] (no locality given; Bombay, India, here designated); lectotype may be in Berlin Museum. 1866 Melaraphe subgranosa Dunker, Verhandlungen der Kaiserlich-Koniglichen zoologisch-botanischen Ge- sellschaft in Wien, Jahrgang 1866, vol. 16, p. 913 (Madras [India]); type may be in Berlin Museum; 1867, Frauenfeld, Reise der Osterreichischen Fre- gatte Novara, Zoologischen Theil, vol. 2, part 3, Mollusken, p. 9, pl. 1, figs. 10 a, b [not fig 10 ¢ which G. millegrana Philippi]. 1887 Litorina leucostica ‘Philippi’ Tryon, Manual of Con- chology, vol. 9, p. 299 [error for L. leucosticta Phi- lippi, 1847]. Types—Philippi's figure of leucosticta (the lec- totype figure) depicts a young specimen, a phe- notype apparantly quite common in India and Ceylon, although this particular oval white spot- ted form has not been noted in collections from elsewhere. Proportions of the subspecies G. leu- costicta biangulata are quite different. The lecto- type figures of Frauenfeld of subgranosa Dunker, show the species in its mature form. Records—INDIA: Bandra, N. of Bombay; Bombay; Goa; Cape Comorin; Tuticorin; Mandapam Camp. CEYLON: Point de Galle; Columbo (all USNM), Nodilittorina leucosticta subspecies biangulata (von Martens, 1897) (Pl. 384, figs. 6-13) Range—East Indies and [?] western Melanesia. Remarks—Littorina leucosticta biangulata von Martens differs from the nominate subspecies in having typically only two strong spiral sculptural cords on the body whorl. Other sculpture is much suppressed. Shells of this subspecies also tend to be more high spired and therefore the tendency for the spire to be less than half the length of the shell is somewhat reduced. Shells reach 12.4 mm. (about .5 inch) in length; average obesity about .62 (29 specimens range from .56-.65). In the representatives of populations available to me for study, the conservatism in spiral sculpture exhib- ited by the type is shown in only a few speci- mens—most develop 3 moderately strong spiral cords bearing subdued white nodules, more like leucosticta s.s., but with the above-mentioned narrower profile (see illustrations, also remarks under leucosticta leucosticta). Habitat—On intertidal rocks. Measurements (mm.)— length width no. whorls locality 12.4 7.0 6 Touho, New Caledonia 11.0 7.1 6 Holotype: Benkulen Sumatra 10.4 6.8 5+ Jesselton, No. Borneo. 10.1 6.0 54+ Touho, New Caledonia 9.7 6.0 4+ Touho, New Caledonia 8.6 ee, 5+ Touho, New Caledonia 7.8 D0 6 Jesselton, No. Borneo 7.1 4.4 6 Corregidor Id., Luzon, Philippines 6.7 4.2 5+ Pulau Nias, Sumatra 5.7 3.6 6 Pulau Nias, Sumatra Synonymy— 1897 Littorina biangulata von Martens, in Max Weber, Zoo- 2 = logische Ergebnisse einer Reise in Niederlandisch [05-403] 500 Granulilittorina Ost-Indien, vol. 4, part 1, p. 209, pl. 9, fig 26 (Benkulen [Bengkulu, SW Sumatra]); holotype in Zoologisch Museum Amsterdam, 11 x 7.1 mm. Records—PHILIPPINES: West tip Corregidor Id. Luzon (ANSP, USNM); Santo Domingo, Batan Id., Batanes Group (USNM). EAST INDIES: Jesselton, N. Borneo; Pulau Nias, (both USNM); Bengkulu, both S.W. Sumatra (Von Martens, 1897; ZMA): Kahatola Id., S. Loloda Ids. Halmahera, Moluc- cas; Pelabuhan Ratu, Preager, Java (both USNM); NEW CALEDONIA: mouth Hienghene R., Hienghene area; 18 km. N. of Touho, N.W. New Caledonia (both USNM). Nodilittorina leucosticta subspecies feejeensis (Reeve, 1857) (PI. 384, figs. 14-19) Range—Pacific Islands, exclusive of Western Melanesia and Hawaii. Remarks—According to available records Gran- ulilittorina leucosticta feejeensis replaces leuco- sticta s.s. and l. biangulata in the Pacific Islands. It usually has a smaller, more compact shell and spiral sculpture and axial color striping are often strongly expressed, although from the present appearance of Reeve’s type, this would not be expected. The type of L. feejeensis Reeve has apparently undergone some corrosion of sculp- ture in over 100 years in the BM(NH) collection. Shells reach 12.7 mm. (about .5 inch) in length; average obesity about .64 (23 specimens range from .61-.72). Most strikingly colorful popula- tions appear to occur in Samoa and Tonga where zigzag dark axial markings stand out against the lighter ground color of the shell (see pl. 384, figs. 18, 19). The general shape of the shell and charac- ter of sculpture and coloration clearly relate this species most closely to leucosticta and I. bian- gulata. Habitat—Shore rocks. Measurements (mm.)— length width no. whorls locality ay 7.8 5+ Ofu, Manu’a, Samoa 11.0 7.5 5 Ofu, Manu’a, Samoa 10.3 6.4 5 Tongatapu 9.4 6.0 5 Lifu, Loyalties 8.7 5.4 6 Arue, Tahiti 8.2 5.7 5 Niuafou 7.6 5.5 4 Makatea, Tuamotus 7.6 5.1 5 Kermadecs 6.5 4.5 5 Niuafou 5.6 4.0 4 Tutuila, Samoa Synonymy— 1857 Littorina feejeensis Reeve, Conchologia Iconica, vol. 10, Littorina, pl. 15, figs 82 a, b. (Feejee [sic] ids.); lectotvpe BM(NH) 1968319: 7.1 x 5 mm. 1871 Litorina vitiensis ‘Reeve’ von Martens, Donum Bis- marckianum, p. 40; emendation of L. feejeensis Reeve, 1857. 1871 Litorina (Melaraphe) vitensis Dunker, Malakozoo- logische Blatter, vol. 18, p. 150 (Hab. ad insulas Vitenses). Joseph Rosewater Littorinidae 1885 Littorina plena var. vitiensis ‘Dunker’ Nevill, Hand-List of Mollusca in the Indian Museum, Calcutta, Part 2, p. 139 (Tahiti); emendation of L. vitensis Dunker, 1871. 1885 Littorina miliaris var. fijiensis ‘Reeve’ Nevill, ibid., p. 154 (Tahiti); emendation of L. feejeensis Reeve, 1857. 1951 Nodilittorina miliaris ‘Quoy and Gaimard’ Habe, Illus- trated Catalogue of Japanese Shells, vol. 1, no. 14, p. 92, pl. 14, fig 5; not N. miliaris (Quoy and Gaimard) which is Eastern Atlantic species and not from As- cension Id., Pacific = Ponape, Caroline Ids. Records—(All USNM except where noted). BONIN IS- LANDS: Port Lloyd, Chichi Jima; Ani Jima. MELANESIA: Vanikoro, Santa Cruz Group (AMS); Lifu, Loyalties. FIJI: W. side Koro Levu Id. Taveuni. KERMADECS: Raoul, (AMS). SAMOA: Pago Pago, Tutuila; Tafuna, Tutuila; Ofu, Manu’a Group. TONGA: Niuafou; Niutoua, fringing reef, Tongatapu; Lualea reef, nr. Fatuma, Tongatapu. SOCIETIES: Puputeai, nr. Mt. Taharaa, Arue, Tahiti; Mt. Taharaa, Dist. of Mahina, Tahiti; N. coast of Tahiti. TUAMOTUS: Makatea (USNM, ANSP); N. of Tamao Hbr., Makatea; Maiai Id., Tikahau. Nodilittorina exigua (Dunker, 1860) (PI. 386, figs. 1-6) Range—Coasts of China and Southern Japan. Remarks—Although G. exigua is apparently closely related to G. millegrana Philippi, an ex- amination of representatives of Japanese and China coast populations reveals a species that is generally more strongly sculptured, lacks the color striping, has a stronger siphonal trough and more deeply excavated columella than mill- egrana. Habe (1951) synonymized exigua under Nodilittorina granularis (Gray) but it has been demonstrated that granularis is a synonym of the Atlantic species, N. miliaris (Quoy and Gaimard). (see remarks under millegrana). Habitat—Shore rocks. Description—Shell reaching 12.5 mm. (about 0.5 inch) in length, high-spired globose to turbi- nate; average obesity about .68 (22 specimens range from .64 to .72); moderately thick in struc- ture, imperforate; spiral sculpture usually prom- inent and consisting of closely spaced raised spiral cords which are alternately granulose and smooth; granulose cords considerably more con- spicuous; axial sculpture limited to occasionally coarse oblique lines of growth. External color fairly constant, usually a rather dirty yellowish to grayish white. Aperture medium to dark reddish brown; inner edge of outer lip often lighter colored with spots of brown marking sculptural furrows; with a broad white band revolving in- ward from near anterior junction of outer lip and columella; columella usually lighter than aper- ture, occasionally as dark. Base only moderately November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 501 Plate 386. Figs. 1-6. Nodilittorina (Granulilittorina) exigua (Dunker) Figs. 1,2. Lectotype figures of Litorina exigua Dunker, from Japan (from Dunker, 1861, Mollusca Japonica, pl. 2, fig. 3). Figs. 3,4. Matsu Shima, Sea of Japan (USNM 601593, 9.8 » 6.8 mm.). Figs. 5,6. Imaizumi, Kagoshima Bay, Japan (USNM 363708, 8.9 x 6 mm.). Figs. 7-12. Nodilittorina (Graulilittorina) picta (Philippi). Figs. 7,8. Lectotype of Littorina picta Philippi (BM(NH) 1968324, 9.8 x 6 mm.). flattened; periphery not carinate. Whorls 4-5, rounded. Spire less than half the length of shell, produced at an angle of from 68-78°. Aperture oval; outer lip moderately thick, usually wrinkled at edge; inner lip (columella) moderately well developed, rather deeply excavated anteriorly, with a small denticular bump usually evident one third the distance antero-posteriorly; having a short but pronounced anterior siphonal trough; with a flattened crescent-shaped area on base adjacent to columella callous. Suture moderately well impressed. Nuclear whorls light brown but worn in all specimens examined. Operculum pau- cispiral, dark brown and oval. Details of animal and reproduction unknown. ‘XS 2 ae >| 5 Figs. 9,10. Smooth form of N. picta; Mokuoloe Id., Oahu, Hawaii (USNM 346407, 7.2 x 4.7 mm.). Figs. 11,12. Nodulose form of N. picta; same locality as smooth form (USNM 346411, 8.5 « 5.6 mm.). Figs. 13-18. Nodilittorina (Granulilittorina) cinerea (Pease). Figs. 13,14. Lectotype of Littorina cinerea Pease [Mar- quesas Islands] (ANSP 18811, 7.9 x 5.1 mm.). Figs. 15,16. Moderately sculptured form, Atuona Bay, Hivaoa Id., Marquesas (ANSP 155486, 8.9 x 5.7 mm.). Figs. 17,18. Fatu Hiva, Marquesas (USNM 697101, 6.9 x 4.6 mm.). Figs. 19,20. Holotype of Littorina iwakiana Nomura and Hatai, from Tanagura Miocene, Japan (Saito Ho-on Kai Museum, Sendai, Japan, Reg. no. 6895, 3.8 x 3.2 mm.). ___ 160° I. | EXIGUA | BONIN <> 1S, 4 lwo Jima a eye| MARIANA ? is | Soipon ° Guar , tok « Yop , 1 iti Eniweto ei Moe. | : . fee wy + Paha CAROLINE “Gas Tes Plate 387. Geographical distribution of Nodilittorina (Granu- lilittorina) exigua (Dunker), [05-405 ] 502 Granulilittorina Measurements (mm.)— length width no. whorls locality 12.4 8.5 4+ Matsushima, Japan 11.2 7.6 4 Matsushima, Japan 10.8 Lio 4+ Komoi, Awaji, Japan 10.2 7.1 4 Matsushima, Japan 9.8 6.3 4 Peiyushan Id., China 9.1 6.2 4 Komoi, Awaji, Japan 8.8 6.3 4 Imaizumi, Japan 8.4 5.9 4 Shirahama, Japan 40 a 4 Hakodate, Japan 6.9 4.7 3+ Takami, Japan Synonymy— — Littorina granularis ‘Gray’ of Authors, non Gray, 1839; Gray's holotype, BM(NH) 87.4.26.9 is the Eastern Atlantic species, Nodilittorina miliaris (Quoy and Gaimard), 1860 Litorina exigua Dunker, Malakozoologische Blatter (1859), vol. 6, p. 226 (Japan); 1861, Dunker, Mollusca Japonica, p. 13, pl. 2, fig. 3 [lectotype figure]. Joseph Rosewater Littorinidae Types—The type specimen of L. exigua Dunker is probably in the Berlin Museum al- though it has been impossible to locate it with certainty during the present study. The specimen figured by Dunker in the year following the original description is here considered the lecto- type (see pl. 386, figs. 1, 2). Records—(All from USNM except where noted). CHINA: Sidesaddle Island (Lu-hua_ shan), Chekiang Province; Peiyushan Id.; Spider Island, Fukien Province; Big Wave Bay, Hong Kong; Cape D’ Aguilar, Hong Kong. JAPAN: Hako- date, Hokkaido; Saigo, Dogo Ids. Oki Group; Matsu-Shima; Shirahama, Wakayama Province, Honshu; Toshima, Tanabe Bay, Honshu; Awaji-shima; Kii, Honshu (AMS); Tokyo Bay (ANSP); Imaizumi, Kagoshima Bay, Kyushu (USNM, ANSP); Tosa Shikoku; Hachijo Island, 275 miles S. of Tokyo (both ANSP). [05-406] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 503 Nodilittorina picta (Philippi, 1846) (Pl. 386, figs. 7-12) Range—Hawaiian Islands. Remarks—The “Painted Littorine” Granulilit- torina picta is closely related to G. millegrana, G. exigua, and apparently also to G. cinerea Pease. The four species all possess similar variation in sculpture. The relationship with millegrana and exigua was pointed out by Struhsaker (1968a) who suggested that picta is the only one which develops a smooth shell as well as a granulose one [some populations of millegrana develop smooth shells also; see pl. 386, fig. 7,8, 9, 10]. The possible selective processes causing the extensive shell variation in picta are well discussed by Struh- saker (ibid.). It is probable that morphology of most wild populations is under the positive con- trol of natural selection there being few if any mutations which survive for very long in nature without some selective value. [The survival of so many natural populations is coming under the aegis of man, that the beneficial “weeding out” effect of unsuitable elements by natural selection may be considerably lessened. One wonders what eventual effect this will have on all species in- volved, including Homo sapiens.] It is, therefore, most likely, as pointed out by Struhsaker that each of the shell forms of picta, from highly granulose to smooth, has survival value under particular ecological conditions, i.e. smoother forms survive best in wave-swept areas, and sculptured forms occur in sheltered localities. The morphological differences are apparently ge- netically linked, but survival of the resultant phenotype is under the control of environmental selection. The subspecies name L. picta marmorata Phi- lippi has generally been used for one of the var- iations of picta in Hawaii. The name marmorata was not validly introduced by Philippi in connec- tion with the original description of picta (see synonymy) but was mentioned and figured in the “Abbildungen” the following year with the add- ed locality “provincia Ilocos borealis insulae Lucon” [Philippines]. Both figure and locality indicate that L. picta marmorata is a synonym of G. millegrana. It is likely that Philippi, himself, did not clearly discriminate between the popula- tions of picta from Hawaii and some of the forms of millegrana occurring in the Philippines. They are indeed similar and this has given rise to considerable confusion, wherein extra Hawaiian Granulilittorina are called “picta.”’ As the mem- bers of this subgenus all appear to be fairly closely related it is perhaps a matter of personal preference whether one considers them separate species or geographic subspecies. In the case of picta and its allies it is here considered less confusing to consider them separate species. In likening picta to neritoides, Philippi indica- ted that he was describing a rather smooth little shell, and the three syntypes of picta in the British Museum (NH) reinforce this view. In my experience, the name “marmorata”™ has been used for the more highly sculptured forms, although certainly the figure given by Philippi does not show such a shell. It would appear that the local interpretations of these names have come about with usage. However, when the situation is ex- amined closely it is found that picta is the only valid name for this species; marmorata, as shown above, not only is a synonym of another species but is preoccupied by L. marmorata Pfeiffer, 1839 [= L. saxatilis Olivi]. Tinker’s (1952) sug- gestion that picta is considered by some to be a synonym of L. planaxis (eastern Pacific) is under- standable. Certain populations especially of young planaxis show similarities to picta in color variation. They are, however, in distinct generic groups, as well as being separate species (see List of Taxa). The connection between L. planaxis and picta probably dates from Tryon’s (1887) synony- my. Habitat—Shore rocks above high tide line (see Struhsaker, 1968a). Description—Shell reaching 12.9 mm. (about 0.5 inch) in length; rather conic to turbinate in shape; average obesity about .64 (20 specimens range from .59-.69); moderately thick in structure, imperforate; surface often smooth, or only micro- scopically spirally striate; but often bearing raised, rather strong and granulose spiral cords on spire and body whorls [granular sculpture most similar to that in G. millegrana, but whorls shouldered as in G. leucosticta and its subspe- cies]. Axial sculpture consists of faint to rather coarse, irregular lines of growth. External color very variable: generally consisting of grayish to yellowish white ground color “painted” with dark-brown markings, a common pattern on body whorl being the center of whorl dark, with area above and below divided into light colored sec- tions by wavy brown lines; The latter not at all constant and brown painting may be minimized or maximized; more highly sculptured shells ap- pear darker overall. Aperture medium to dark brown and may be brown and white mottled as is [05-407] 504 Granulilittorina exterior; with an often diffuse or interrupted white band revolving inward from near anterior junction of outer lip and columella; columella usually light tan; inner edge of outer lip white or with a few white spots. Base slightly flattened; periphery occasionally with a weak to moderate keel. Whorls 5-6, not very well-rounded, body whorl distinctly flat-sided. Relative lengths of spire and aperture about co-equal, one sometimes exceeding the other apparently indiscriminately; spire produced at an angle of from about 60-65°. Aperture oval; outer lip moderately thick; inner lip (columella) only moderately developed, witha flattened crescent shaped area on base adjacent to columella callus. Suture well impressed. Sculp- ture varying from virtually smooth to markedly spirally striate with coarse granulations. Nuclear whorls light brown anteriorly, darker brown pos- teriorly on each whorl, smooth and unsculptured; in shells which will be granulosely sculptured, this begins after about 21/2 nuclear whorls; in shells which never develop coarse sculpture, only low spiral striae begin after nuclear whorls. Oper- culum paucispiral, brown, oval. Radula littori- nid, 2-1-1-1-2, central tooth rather narrow. Details of animal are from Whipple (1965): In males base of penis is red as is testicular duct; there is an enlargement at base of penis partially separated from a penial gland containing an ac- cessory flagellum; sperm groove runs along pos- terior edge, deeply folded. Reproduction ovipa- rous, producing a sculptured pelagic capsule about 180m in diameter, with single egg about 75u in diameter. According to Struhsaker (1968b) a swimming veliger hatches in about 3 days and the larva settles and metamorphoses about 21 days after spawning. Measurements (mm.)— length width no. whorls locality 12.9 8.0 5+ Hawaiian Ids. 12.7 8.0 6 Waikiki, Oahu 11.8 7.9 5 Waikiki, Oahu te 6.6 ay btwn Waipio and Pearl City, Oahu Joseph Rosewater Littorinidae 10.9 6.8 5 Diamond Head, Oahu 10.0 6.8 5 Launiupoku, Maui 9.9 6.1 5 btwn Waipio and Pearl City, Oahu 9.6 6.1 5 Paia Beach, Maui 9.1 5.9 6 Wainini, Kauai Synonymy— 1846 Littorina picta Philippi, Proceedings of the Zoological Society of London, for 1845, part 13, p. 139 (Hawaii); lectotype in BM(NH) 1968324: 9.8 x 6.0 mm.; 1847, as Litorina picta Philippi, Abbildungen und Besch- reibungen Conchylien, vol. 2, Litorina p. 166 [not L. picta marmorata, ibid., p. 167, pl. 3, fig 26 (from Luzon, Philippines) = G. millegrana Philippi]. 1857 Littorina picta Philippi, in Reeve, Conchologia Iconica, vol. 10, Littorina pl. 15, figs 80 a, b, 81. 1887 Littorina planaxis ‘Nuttall’ Tryon, Manual of Concholo- gy vol. 9, p. 248, pl. 44, fig 57 [only in part; includes L. picta Philippi as synonym]; not L. planaxis ‘Nut- tall’ Philippi, 1847, an eastern Pacific species. Types—Three syntypes of Littorina picta Phi- lippi were found in the British Museum (NH) 1968324. Of these, the one most nearly approxi- mating the original measurements is here desig- nated lectotype. Original measurements given by Philippi were: “Altit. 4, diam. 3 lin.” which yields the following millimeter measurements based on a “German Line” equaling 2.18 mm. (Rehder, 1945): length 8.7, width 6.6 mm. The actual measurements of the designated lectotype are 9.8 x 6.0 mm., fairly close to the original, and here considered within the limits of a possible measurement error. (see pl. 386, figs. 7, 8) Records—HAWAIIAN ISLANDS: Laysan Island : La Pe- rouse Rock, French Frigate Shoals; Nihoa Island (all BPBM). KAUAI ISLAND: Nawiliwili (BPBM): Koloa (ANSP); Haena (BPBM). OAHU: Kahuku Point (BPBM); Laie (ANSP, MCZ); Kahana; Mokolii Island, Kaneohe Bay; Mokuoloe Island; Moku Manu; Kailua; Manana (Rabbit Island) (all BPBM): Diamond Head; Waikiki (both ANSP, MCZ, BPBM); Pearl Harbor; Barber’s Point (both BPBM): off Puu Mailiilii (MCZ);: E. of Waianae (ANSP); Makua (MCZ); Haleiwa (ANSP); Waimea (BPBM). MOLOKAI: Pauwalu; Papohaku (both BPBM); Moomomi (BPBM, ANSP); . MAUI: Lahaina (ANSP). KAHOOLAWE ISLAND: (ANSP. HAWAII: Hilo (ANSP, BPBM, MCZ); Kalapana; Waiahukini, Kau; Hoopuloa (all BPBM); Refuge City, Honaunau Bay, Kona (ANSP); Keauhou (BPBM); Kahaluu (ANSP); Kaulana, Kau; Tawai, Kalalaa, at stream outlet (both BPBM). [05-408 ] November 30, 1970 Nodilittorina cinerea (Pease, 1869) (Pl. 386, figs. 13-18) Range—Marquesas Islands. Remarks—N. (Granulilittorina) cinerea is closely related to N. picta of Hawaii and shows much of the same sort of variation from smooth to granulose sculpture. A color pattern is also evi- dent in occasional specimens of cinerea, but never to the same degree as in picta. As in the case of picta, cinerea is also quite closely related to the other species of the subgenus Granulilittorina, and some of its forms are very close to N. mil- legrana. Major distinguishing features are (1) the gray color as indicated by Pease’s name “‘ci- nerea’, which never reaches the marbled colora- tion of picta, (2) the whorls which may become relatively broadly shouldered, and (3) the degree of sculpture, ranging from almost smooth to fairly granulose spiral cords. According to available records, this species is endemic in the Marquesas Islands. Habitat—In splash zone, above high tide line, in cracks in lava, crawling on rock surface at night (personal communication, H. A. Rehder; from observations made at Nuku Hiva, Marquesas Is- lands, 1967). Description—Shell reaching 9.0 mm. (about /3 inch) in length; turbinate to shouldered-turbinate in shape; average obesity about .65 (20 specimens range from .58-.71); only moderately thick in structure; imperforate; sculpture of low to rather distinct spiral cords; cords smooth or granulose; whorls varying from only slightly to rather dis- tinctly shouldered; axial sculpture consisting of faint to coarse growth lines and furrows; occa- sionally specimens have a deep, ragged axial mark of growth interruption. External ground color somewhat variable from grayish white to distinct bluish gray, the latter especially in the young; occasional patterning consists of irregular darker gray to brown zigzag lines and flammules. Aperture medium reddish brown to dark brown with a white band revolving inward from near anterior junction of outer lip and columella; colu- mella usually colored about as aperture or slightly lighter; inner edge of outer lip white. Base some- what flattened; periphery not keeled, whorls shouldered for about #/s the distance anterior to suture. Whorls about 4-6, not too well rounded but rather tending to be straight-sided. Relative INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 505 lengths of spire and aperture about coequal or spire slightly shorter. Spire produced at an angle of from about 60-67°. Aperture widely oval; outer lip moderately thick; inner lip (columella) moder- ately strong, with an often well depressed cres- cent shaped area on base adjacent to columella callous; the depression rarely approaching an umbilical chink. Suture well impressed. Sculp- ture varying from specimens with hardly raised spiral cords to those exhibiting distinct well raised cords with granulations on all whorls. Nuclear whorls light brown anteriorly, dark brown posteriorly smooth and unsculptured; nu- cleus about 2 whorls in extent; first postnuclear whorl smooth, subsequent whorls with spiral sculpture; overall microscopic spiral sculpture consisting of fine wavy spiral threads. Operculum paucispiral light brown, oval. Radula littorinid, 2-1-1-1-2, central tooth rather narrow. Animal similar to other Nodilittorina. Nothing is known concerning reproduction and _ larval development; probably similar to G. picta. Measurements (mm.)— length width no. whorls locality 9.0 5.8 5+ Hiva Oa, Marquesas 8.7 5.3 4+ Hiva Oa, Marquesas 8.5 5.6 5+ Hiva Oa, Marquesas 8.2 4.8 6+ Hiva Oa, Marquesas 8.1 5.2 4+ Hiva Oa, Marquesas 7.9 5.1 6 Lectotype, Marquesas 13 bd. 4+ Fatu Hiva, Marquesas 7.0 AT 3+ Fatu Hiva, Marquesas 6.8 4.6 4+ Fatu Hiva, Marquesas 6.4 4.3 4+ paralectotype, Marquesas Synonymy— 1869 Littorina cinerea Pease, American Journal of Concholo- gy, vol. 5, part 2, p. 78, pl. 8, fig. 14 (Marquesas Islands); lectotype ANSP 18811: 7.9 x 5.1 mm. Types—tThere are three syntypes of Littorina cinerea Pease in the ANSP. Of these, one, most resembling the figure associated with the original description, is here designated lectotype: ANSP 18811 (pl. 386, figs. 13, 14). Its measurements: length 8.1, width 5.2, are fairly close to the originals given by Pease: 7 x 5.5 mm. The lecto- type and two paralectotypes fairly well fulfill Pease’s descriptive comments: “... . transversely granosely ridged or smooth.... whitish, apex blackish ..... ° Records—MARQUESAS ISLANDS: Nuku Hiva (ANSP; USNM);: Ua Pou; Tahuata (both USNM):; Ua Huka (ANSP); Hiva Oa (ANSP, USNM); Fatu Hiva (USNM). cinereous or [05-409] 506 Granulilittorina Joseph Rosewater Littorinidae Nodilittorina iwakiana (Nomura and Hatai, 1936) (Pl. 386, figs. 19, 20) Range—Miocene, Japan. Remarks—The tiny species described by No- mura and Hatai appears to show closest affinity with the members of the subgenus Granulilit- torina. Details of sculpture given by the authors suggest that this fossil may be related to G. millegrana Philippi. Synonymy— 1936 Littorina iwakiana Nomura and Hatai, Saito Ho-on Kai Museum Research Bulletin, no. 10, p. 144, pl. 16, figs 16a, b. (Tanagura Miocene, Okada, Japan); unique holotype in Saito Ho-on Kai Museum, Sendae, Japan, Reg. No. 6895; 3.8 x 3.2 mm. Published by THE DEPARTMENT OF MOLLUSKS Delaware Museum of Natural History Greenville, Delaware 19807, U.S.A. [05-410] January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Tectariinae 507 WILL IAM H. |} SECTIONAI DIVISIO 4 THE FAMILY LITTORINIDAE IN THE INDO-PACIFIC Part Il. The Subfamilies Tectariinae and Echinininae by JOSEPH ROSEWATER Division of Mollusks National Museum of Natural History Washington, D.C. 20560 U.S.A. Abstract The classification of tropical Indo-Pacific Tectariinae and Echinininae (Mollusca: Meso- gastropoda: Littorinidae) has been revised. Seven Recent and three Tertiary fossil species belonging to three generic or subgeneric groups are rede- scribed and figured. Complete systematic synon- ymies are given, together with discussions of relationships, biology and zoogeography. A list is given of world-wide members of Tectariinae and Echinininae. Introduction In Part I of Indo-Pacific Littorinidae the spe- cies belonging to the subfamily Littorininae were reviewed by Rosewater (1970). The present sec- tion covers the subfamilies Tectariinae and Ech- inininae. As yet not covered are such groups as Bembicium and its allies and Cremnoconchus, although they both occur in the Indo-Pacific re- gion. They are here considered to constitute at least subfamily groups, if not separate families (see Bibliography in Rosewater, ibid., p. 427: Anderson, D. T., 1960, and also Anderson, H., 1958; also see Prashad, 1925 and Kesteven, 1903). The many Antarctic littorinids were excluded from Part I of this study since actually they are out of the geographic area of the tropical Indo- Pacific (see Powell, 1951 and Dell, 1964). The Littorininae, Tectariinae and Echinininae differ in habitat preference and in morphology in a number of aspects. Echinininae tend generally to live rather high on the shore. Tectariinae are next and Littorininae usually are nearest to the sea, although considerable variation exists, the genus Nodilittorina having some high-living spe- cies. The differentiation of the three subfamilies also is based on the following morphological grounds. In Littorininae the radula is of the gen- eralized littorinid type with some narrowing of the central tooth in Nodilittorina; opercula are paucispiral; penises are adorned with a few peni- al glands but otherwise are not particularly com- plicated. In the Tectariinae the lateral radula tooth is partitioned and appears thickened and differently oriented than in either Littorininae or Echin- ininae; the opercula are mesospiral (see below); penises are abundantly supplied with glands and are also papillose on non-glandular surfaces. The shells in both Littorininae and Tectariinae usual- ly are imperforate although the subgenus Cen- chritis is an exception in the latter subfamily. In Echinininae the radula is not unusual, the subgenus Tectininus excepted, where reduction has taken place; opercula are multispiral; shells are umbilicate, usually; penises are supplied with a number of penial glands near the base. In the possession of an umbilicus Cenchritis could be considered to form a bridge between Tectarius and Echininus. In general, however, Tectarius appears to be more closely related to Littorina in its conservative characters, while Echininus may be considered a specialized group perhaps evolv- ing toward a land environment. These three subfamily groups provide a convenient and ap- parently natural framework upon which to ar- range the contained genera and species (see illus- [05-441] 508 Joseph Rosewater Littorinidae Plate 388. Subfamilies Tectariinae and Echinininae (explanation on opposite page; all figures about natural size) [05-442] January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Tectariinae 509 trations of these morphological differences in Rosewater, 1970, and in this paper). Most Tectarius s.s.and Echininus s.s. are living today only in the East Indian area. With the exception of Tectarius grandinatus whose range extends eastward to Polynesia, all other species are inhabitants of the raised, weathered coral reef shorelines found in the Western Pacific Arc (per- sonal observations, 1970). It can only be assumed that this niche provides the requirements essen- tial for the existence of these species‘as they are to be found nowhere else. The fossil record provides very few clues to the origin of these groups. There are only three Terti- ary fossil species described from the Indo-Pacific and these probably belong in three separate ge- neric taxa. The oldest of these, T. songoense Martin, from the Upper Eocene of Java, probably represents nearly the earliest appearance of Tec- tarius. As mentioned in Part I, littorinid fossils are exceedingly difficult to separate from Trochi- dae and Turbinidae, and this is no less true for Tectarius and Echininus. Probably, however, these groups made their appearance in the early Tertiary within the region where they have devel- oped, and with the exceptions of Echininus (Tec- tininus) nodulosus and Tectarius (Cenchritis) muricatus, both of the western Atlantic, they have remained there. Opercula The opercula of Indo-Pacific Littorinidae re- quire special comment (see pl. 389). All are made up of conchiolin, and those of members of the subfamily Littorininae, including Littorina, its subgenera, and Nodilittorina are usually pauci- spiral and rather oval in shape with the nucleus at the side and nearer one end (oligogyrous spiral type of Fretter, et al., 1962, pp. 79,80). In the Echinininae, the basic plan of the operculum differs from that of other littorines. It is the type Plate 389. Opercula,of Littorinidae and Trochidae. Fig. A. Paucispiral operculum of Littorina (Littorinopsis) scabra (Linné) from Mokuoloe Id., Kaneohe Bay, Oahu (USNM 339388). Fig. B. Mesospiral operculum of Tectarius rusticus (Philip- pi) from Troughton Chain, northern Western Australia (WAM 1787-69); Fig. C. Multispiral operculum of Echininus cumingi (Phi- lippi) from near Davao City, Mindanao, Philippines (WAM 1566-70); Fig. D. Multispiral operculum of Trochus niloticus Linne, from Makuluva, Viti Levu, Fiji (USNM 531827). Lines under each figure represent 5 mm.; stippled areas are thickened and dark-brown in color; non-stippled areas are ight horn color and transparent. of operculum which is called in other groups, such as Trochidae, a multispiral operculum (polygyrous spiral type of Fretter, et al., ibid.) although not so extreme as that figured by Fetter (ibid., p. 80, fig. 43A; also see our pl. 389, fig. D.). The operculum is circular in outline and moder- ate to small in size. The nucleus is decidedly central in location and growth proceeds outward from the center in multiple, fairly evenly spaced gyrations (pl. 389, fig. D). The operculum in Tec- tarininae (fig. B) is intermediate in form between Explanation to plate 388 (opposite page) Figs. 1,2. Tectarius grandinatus (Gmelin) from Palmerston Atoll, Cook Islands (USNM 685165) Figs. 3,4. Tectarius tectumpersicum (Linné). Fig. 3, from Stirling Isle, Treasury Ids., Solomon Islands (USNM 600370); Fig. 4, from “East Indies” (USNM 131450). Figs. 5-7. Tectarius pagodus (Linné). Fig. 5, from “East Indies” (USNM_ 18966); Fig. 6, from the Philippines (USNM): Fig. 7, a young specimen from Polillo, Philip- pines (USNM 311141). Figs. 8,9. Tectarius rusticus (Philippi) from Buccaneer Archipelago, Western Australia (USNM 684713). Figs. 10,11 Tectarius rugosus (Wood). Fig. 10, from Pacific (USNM 304587); Fig. 11, from Davao Bay, Mindanao, Philippines (USNM 654034). Figs. 12,13. Echininus cumingi cumingi (Philippi) from Hervey Ids., Cook Ids. (USNM 42452). Figs. 14,15. Echininus cumingi spinulosus (Philippi) from Kadena Circle, Okinawa, Ryukyu Ids. (USNM 664658). Figs. 16,17. Tectarius (Cenchritis) muricatus (Linne), from Boca de Camarioca, Matanzas, Cuba (USNM 599944). Figs. 18-20. Echininus (Tectininus) nodulosus (Pfeiffer). Fig. 18, from Hog Island, Bahamas (USNM 603911); Figs. 19, 20, from Mujeres Harbor, Quintana Roo, Mexico (USNM 662308). [05-443] 510 Joseph Rosewater Littorinidae Littorininae and Echinininae, being large and rather rounded in outline, with a slightly acentric nucleus and having a number of gyrations more than the paucispiral type but less in number than the multispiral type. The Tectariine opercular type is here termed the mesospiral or mesogyrous spiral type. The presence of the three opercular types in Littorinidae may be considered to have evolutionary significance, and possibly is related to selection for a better aperture sealing mecha- nism in animals which have considerable vertical distribution on the shore line: in order, proceed- ing from low toward higher shore habitats— Littorininae, Tectariinae, Echinininae. Reproduction To my knowledge nothing is known concern- ing reproduction in either Tectariinae or Echin- ininae with the exception of Tectarius (Cen- chritis) muricatus (Linné) which produces a pe- lagic capsule (see Lebour, 1945, and Lewis, 1960, references in Rosewater, 1970 p. 05-276). Field and laboratory studies are needed to discover details of the life histories of the remaining spe- cies. However it is likely that most of these snails also produce eggs encased in pelagic capsules which undergo development in the sea. Abbott (1954) noted that Lebour (ibid.) stated that some of the Bermuda littorinids that live above high tide line migrate to the water to spawn. It is suspected that this also is the case with many of the Indo-Pacific species. Acknowledgments The persons and institutions acknowledged in Part I of this study (see Rosewater, Indo-Pacific Mollusca, vol. 2, no. 11, p. 425) also are thanked here. In addition, I acknowledge the following for their help in making possible the examination in the field of most of the species of Tectarius and Echininus during the National Geographic Socie- tvy—Mariel King Memorial Expedition to the Mo- luecas Islands, Indonesia, May to July 1970: the late Mariel King, Mrs. Grace King, T. H. Richert, C. Beal, C. M. Burgess, B. R. Wilson, and the National Geographic Society. The Government of Indonesia graciously provided clearance for the vessel Pele to work in the Moluccas Islands. Mr. Kasim Moosa and Mr. Sukarno, both of the Insti- tute for Marine Research, Djakarta, accompanied the expedition and provided assistance of many kinds. List of Recognized Taxa Below is a list of the Tertiary fossil and Recent species herein recognized as belonging in the subfamilies Tectariinae and Echinininae. The few fossil taxa are preceded by a dagger [f]. Family Littorinidae Gray, 1840 Subfamily Tectariinae, new subfamily GENUS Tectarius Valenciennes, [1832] Subgenus Tectarius Valenciennes, [1832] rugosus (Wood, 1828). Type. Recent, wes- tern Pacific grandinatus (Gmelin, 1791). Recent, Pacific islands pagodus (Linné, 1758). Recent, western Pa- cific tectumpersicum (Linné, 1758). Recent, wes- tern Pacific rusticus (Philippi, 1846). Recent, northern Australia + songoense (K. Martin, 1931). Eocene, Java. Subgenus +Subditotectarius Ladd, 1966 trehderi Ladd, 1966. Type. Miocene, Mar- shall Islands. Subgenus Cenchritis von Martens, 1900 muricatus (Linne, 1758). Type. tropical western Atlantic. Recent, Subfamily Echinininae, new subfamily GENUS Echininus Clench and Abbott, 1942 Subgenus Echininus Clench and Abbott, 1942 cumingi cumingi (Philippi, 1846). Type. Re- cent, western Pacific cumingi spinulosus (Philippi, 1847). Recent, western Pacific + adelaidensis (Cotton, 1947). Pliocene, South Australia, Subgenus Tectininus Clench and Abbott, 1942 nodulosus (Pfeiffer, 1839). Type. Recent, tropical western Atlantic. [05-444] January 15, 1972 Selected Bibliography Abbott, R. T. 1954. Review of the Atlantic Periwinkles, Nodilittorina, Echininus, and Tectarius. Proceedings of the United States National Museum, vol. 103, no. 3328, pp. 449-464. Argenville, A. J. D. d’. 1742. L’histoire naturelle —La Litho- logie et la Conchyliologie —par—M. [A. J. D. d’ Argenville] de la Société Royale des Sciences de Montpellier. Paris. Bruguiére, M. 1792. Encyclopédie Méthodique, Paris, vol. 1, p. 530. Clench, W. J. and R. T. Abbott. 1942. The Genera Tectarius and Echininus in the Western Atlantic. Johnsonia, vol. 1, no. 4, pp. 1-4. Dance, S. P. 1967. Report on the Linnaean Shell Collection. Proceedings of the Linnean Society of London, vol. 178, no. 1, pp. 1-24, 10 pls. Dell, R. K. 1964. Marine Mollusca from Macquarie and Heard Islands. Records of the Dominion Museum, vol. 4, no. 20, pp. 267-301. Deshayes, G. P. 1830. Encyclopédie Méthodique, Paris, vol. 2, p. 184. Dodge, H. 1959. A Historical Review of the Mollusks of Linnaeus. Bulletin of the American Museum of Natural History, vol. 118, Article 5, pp. 211-257. Fretter, V., and A. Graham. 1962. British Prosobranch Mol- luses. Ray Society, London, xvi + 755 pp., 317 figs. Habe, T. 1951. Littorinidae in Japan (1). Illustrated Catalogue of Japanese Shells, no. 14, pp. 87-93. Habe, T. 1961. Coloured Ilustrations of the Shells of Japan (11). Hoikusha Publishing Co., Ltd., Osaka, 183 pp., 66 pls. Habe, T. 1964. Shells of the Western Pacific in Color, Vol. II. Hoikusha Publishing Co., Ltd., Osaka, 233 pp., 66 pls. Kaicher, S. D. 1956. Indo-Pacific Sea Shells. Section 3. Lit- torinacea, ete. Privately Printed, Washington, D.C., 8 pls. and captions. Keen, A. Myra. 1966. Tectarius (Mollusca: Gastropoda): Re- quest for Validation in its Accustomed Sense. Z.N. (S.) 1754. Bulletin of Zoological Nomenclature, vol. 23, part 4, pp. 179-180. INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Tectariinae 51] Kesteven, H. L. 1903. Notes on Prosobranchiata, No. II. Littorinacea. Proceedings of the Linnean Society of New South Wales, 1902, part 4, pp. 620-636. Kira, T. 1959. Coloured Illustrations of the Shells of Japan. Revised Edition, Hoikusha Publishing Co., Ltd., Osaka, ix + 239 pp. Kira, T. 1962. Shells of the Western Pacific in Color. Hoikusha Publishing Co., Ltd., Osaka, 224 pp. Klein, J. T. 1753. Tentamen Methodi Ostracologicae sive Dispositio Naturalis Cochlidum et Concharum, p. 25. Lug- duni Batavorum. Melville, R. V. and W. E. China. 1969. Opinion 871. Tectarius Valenciennes, [1832] (Gastropoda): Validated Under the Plenary Powers. The Bulletin of Zoological Nomenclature, vol. 25, part 6, pp. 214-215. Morch, O. A. L. 1852. Catalogus Conchyliorum Quae Reliquit D. Alphonso D’Aguirra et Gadea Comes de Yoldi; fascicle 1, p. 45. Powell, A. W. B. 1951. Antarctic and Subantarctic Mollusca: Pelecypoda and Gastropoda. Discovery Reports, vol. 26, pp. 47-196, text figs., 6 pls. Prashad, B. 1925. Respiration of Gastropod Mollusks. Pro- ceedings of the Twelfth Pacific Science Congress, pp. 126-143. Rosewater, J. 1970. The Family Littorinidae in the Indo- Pacific. Part I. The Subfamily Littorininae. Indo-Pacific Mollusca, vol. 2, no. 11, pp. 417-506, 64 pls. Rumphius, G. E. 1705. D’Amboinsche Rariteitkamer. Am- sterdam, 340 pp., 60 pls. Sherborn, C. D. and B. B. Woodward. 1901. Bibliographical Notes. XXVIII. The Dates of Humboldt and Bonpland’s “Voyage”. Journal of Botany, June, pp. 1-4. Troschel, F. H. 1856-1863. Das Gebiss der Schnecken, vol. 1, Berlin, pp. vii + 252, pl. 1-20. Watson, R. B. 1886. Report on the Scientific Results of the H.M.S. Challenger, vol. 15, part 42, p. 576. Wenz, W. 1938. Handbuch der Paliozoologie, vol. 6, part 1, lfg. 3, pp. 241-480, figs. 472-1235. Wimmer, August. 1880. Sitzungsberichte der Mathematisch- naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften, Wien, I. Abth. vol. 80, pp. 496, 514. [05-447] 512 Joseph Rosewater Littorinidae Key to the Tectariinae and Echinininae The following key is to the genera and sub- only, since Western Atlantic taxa are not treated in genera of these two subfamilies. It is based upon detail in the present paper. For a key to the the shell and externally observable characters. Littorininae, see vol. 2, no. 11, p. 430 [p. 05-278]. Page numbers are given for Indo-Pacific groups aS etl (iri WCAC ye cus. ea dd a Rhaclea eatacte. Shin al ites dachass lee yh Bh cee een eee 2 Lb: Shell Wor aii TCAte be «scm ted 6 ph ated ate aime we cheer sek De Dh eas Se es ee 3 2a Shell spinose, with partially open spines, shell about as wide as high, operculum maul tigate 166C4pl G00) .< ¢ hae 4 Gnd 4.44 24 4A Ae es Sas Echininus p. 526 2b Shell not spinose, nodulose, higher than wide, operculum not multispiral ... Cenchritis 3a Shell spinose, operculum mesospiral. Tectarius ......0. 5 ccc enna prods oe Shell moculoce: (ose) 3c as swe whee one oe Hab ie 4 PGloe otis Subditotectarius p. 524 3c Shell moderately spinose, operculum multispiral................0.05. Tectininus Plate 390. Type-species of Genera and Subgenera of Tectari- Miocene, Marshall Islands (Holotype, USNM 648342; 2.8 inae (Figs. 1-9) and Echinininae (Figs. 10-15) illustrating x 2.4 mm.). sculpture, arrangement of spines, and presence or absence of Figs. 7-9. Tectarius (Cenchritis) muricatus (Linné); umbilici. Matanzas, Cuba (USNM 599944; 26.1 «x 17.9 mm.). Figs. 10-12. Echininus (Echininus) cumingi (Philippi); Figs. 1-3. Tectarius (Tectarius) rugosus (Wood); Davao Bay, Cook Islands (USNM 42452a; 17.2 « 17.5 mm.). Mindanao, Philippines (USNM 654034; 23.1 x 17.8 mm.). Figs. 13-15. Echininus (Tectininus) nodulosus (Pfeiffer); Figs. 4-6. Tectarius (Subditotectarius) rehderi Ladd; early Cozumel Id., Mexico (USNM 662806; 14.4 x 12.2 mm.). [05-448] January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Tectarius 513 Subfamily Tectariinae, new subfamily Genus Tectarius Valenciennes, [1832] Type: Tectarius rugosus (Wood, 1828) The genus Tectarius sensu lato includes the Tectarius, whose type- species T. rugosus Wood (= T. papillosus ‘La- marck’ of authors) has one of the more conserva- tively sculptured shells of the group. Tectarius sensu stricto is a wholly Indo-Pacific group, hav- ing its present population center in the Western Pacific Arc. The monotypic subgenus Cenchritis von Martens contains only T. (C.) muricatus (Linné), of the tropical western Atlantic (see pl. 388, figs. 16, 17). Subditotectarius Ladd, 1966, is monotypic for the fossil T. (S.) rehderi Ladd, of the Miocene of the Marshall Islands. Only spe- cies belonging to Tectarius s.s. and Subditotec- tarius will be considered here. Tectarius appears more closely related to Lit- torina than to Echininus because of greater simi- larities in morphology, general shell characters, the absence of a truly multispiral operculum, a broader, less modified central radula tooth, and the usual lack of an openly umbilicate shell. nominate subgenus, Subgenus Tectarius sensu stricto Moderately large, pyramidal to turbinate, non-umbilicate littorinids with from rather strongly spinose to nodulose or papillose shells; generally living at or above high tide line. Radula littorinoid, the central tooth somewhat narrowed, the lateral tooth with an embayment and usually developing a medial vertical ridge or partition. In males the penis is large and well-supplied with glands along most of its lateral edge, the remain- der papillose, and with an open but deeply folded seminal duct. Operculum rounded, mesospiral (see Opercula in Introduction). Aperture plicate within; with a columellar swelling or tooth. Synonymy— 1798 Cidaris Roding, Museum Boltenianum, part 2, p. 84; type-species by subsequent designation, Herrmann- sen, 1847: Trochus pagodus Linné; non Cidaris Leske, 1778, nor Swainson, 1840. [1832] Tectarius Valenciennes, Coquilles, in Humboldt and Bonpland. Voyage aux régions équinoxiales du Noveau Continent, Observations de Zoologie, vol. 2, p. 271; type-species by subsequent designation Clench and Abbott, 1942: Trochus coronatus Valenci- ennes [= Tectarius rugosus (Wood]. ICZN Opinion 871. 1839 Pagodus Gray, in Molluscous Animals: The Zoology of Captain Beechey’s Voyage, p. 141; type-species by Monotypy and by absolute tautonymy, Monodonta pagodus Lamarck [= Tectarius pagodus (Linné]. 1840 Pagodella Swainson, A Treatise on Malacology, pp. 207, 219, 221 [refers to Pagodella echinata, nomen nudum| 351; refers to P. major Martini - Chemnitz, pl. 163, figs 1541, 1542 [= Tectarius pagodus (Linné)] and to T. tectumpersicum ibid., fig. 1543, 1544; type-species here designated: Tectarius pagodus (Linné). 1840 Echinella Swainson, ibid., pp. 207, 221, 352; refers to E. granulata Swainson [nomen nudum] and to E. coro- naria, Tableau Encyclopédique et Méthodique, pl. 447, fig 6 [=Mondonta coronaria Lamarck = Tectar- ius grandinatus (Gmelin)]; type-species by monotypy, Tectarius grandinatus (Gmelin) [also see Clench and Abbott, 1942]; not Echinella Bory St. Vincent, 1824. 1846 Fectaria Philippi, Abbildungen und Beschreibungen Conchylien, Vol. 2, Litorina, p. 139; used in combina- tion Fectaria pagodus; error for Tectarius Valencien- nes, 1858 Hamus ‘Klein’ H. & A. Adams, The Genera of Recent Mollusca, vol. 2, p. 656, refers to H. and A. Adams, 1854, vol. 1, p. 315; type-species here designated, Hamus pagodus (Linné) [=Tectarius pagodus (Linné)]; not Hamus ‘Klein? R. B. Watson, 1886 [=Trochidae]. 1899 Echinellopsis Rovereto, Atti della Societa Ligustica di Scienze naturali e geografiche, vol. 10, p. 109; new name for Echinella Swainson, 1840, not Bory St. Vincent, 1824. Nomenclature—Due to similarities between the shells of Tectarius and some of the Trochidae, there has been a tendency for some of the former to be classified with the latter. This problem was discussed by Keen (1966) who recommended that the International Commission on Zoological Nomenclature validate Tectarius with the type- species Tectarius coronatus Valenciennes, [1832], i.e., in its accustomed sense. Her petition was granted in I.C.Z.N. Opinion 871 (Melville and China, 1969). It was assumed in this Opinion that the type-species of Tectarius, T. coronatus Valenciennes, is a synonym of T. grandinatus Gmelin. However, an examination of the type- specimen of coronatus in the Paris Museum shows it to be in actuality T. rugosus Wood, which usually has been erroneously referred to as T. papillosus Lamarck. Another name which sometimes has been as- sociated with Tectarius that has an exceedingly long and complicated history is the genus Ha- mus. It was mentioned originally by Klein (1753) where its use was of course pre-linnaean. Bru- guiére (1792) gave a brief description, referring to Klein, but listed no species. Deshayes (1830) [05-449] 514 Joseph Rosewater Littorinidae declared it “‘a forgotten genus’, indicating that he considered it unrecognizable. Mérch (1852)listed it, this time in the synonymy of Littorina Férus- sac, an invalid introduction (I.C.Z.N., Art. 11(d)). The first valid use of Hamus was not until H. & A. Adams (1858) used it as a senior synonym for Tectarius (see synonymy). It was later used by Wimmer (1880) and Watson (1886), the last being a taxon of Trochidae. I have designated as type- species of Hamus H. and A. Adams, 1858, H. pagodus (Linné) and consider this genus to be an absolute synonym of Tectarius Valenciennes. The use of square brackets surrounding the date for Tectarius Valenciennes, [1832] is recom- mended by the International Code of Zoological Nomenclature in cases where the date of publica- tion of a name has been determined on the basis of external evidence (I.C.Z.N. Recommendation 22.A(3); also see Sherborn and Woodward, 1901; Keen, 1966; and Opinion 871). Tectarius rugosus (Wood, 1828) (Pl. 388, figs. 10, 11) Range—Philippines and Indonesia. Remarks—The shells of well prepared and cleaned specimens of Tectarius rugosus tend to be quite colorful for Littorinidae, with the orange-pink coloration of the last two whorls contrasting with a purplish brown subsutural band. These colors do not show well in all speci- mens, however, and are not very visible in un- cleaned specimens. The three’ large, non- umbilicate common species of the southwest Pa- cific may be distinguished by the number of major spiral rows of spines on the last whorl: 2 in pagodus; 3 in tectumpersicum; and 4 in rugosus. The closely-spaced stubby spines of rugosus also separate it from the other two. These characteris- tics do not of course help to distinguish it from T. grandinatus, but other characters and the Polyne- sian endemicity of the latter are helpful in this case (see Remarks under grandinatus). Habitat—Shore rocks and limestone cliffs 1-2 meters above high tide line (personal observa- tions, Davao, Philippines, 1970). Description—Shell reaching 39.7 mm (about 1?/2 inches) in length, broadly conical in shape, average obesity about .76 (51 specimens range from .68-.82); mature specimens moderately heavily constructed, imperforate, and sculptured on most postnuclear whorls with four, fairly closely-spaced rows of stubby, rounded, often slightly upturned spines. External color generally yellowish white on early whorls, becoming pink- ish orange on penultimate and body whorls; area of most posterior (subsutural) row of spines usual- ly a contrasting purplish brown, and the same dark color may appear in lines and dashes inside outer lip of aperture; aperture tinted lighter pink- ish orange. Base flattened, sculptured spirally with nodulose cords, a larger separate row just below periphery of body whorl. Whorls 6-8, flat- sided excepting spines. Length of spire usually greater than half the length of shell. Spire convex, produced at an angle of from about 60-67°. Aper- ture rounded-squarish; outer lip thickly produced in mature individuals, strongly plicate within; plicae not reaching edge of aperture; outer lip tapering to a thin, crenulate edge; inner lip smooth posteriorly, often stained a deeper orange than rest of aperture, forming a tooth-like bulge anteriorly, near junction with outer lip near base of columella. Suture obscured by anteriormost row of spines of preceding whorl. Primary sculp- tural feature is the four spiral rows of spines. Spines not particularly aligned axially, although anteriormost 2 rows more so than others; from 17-23 spines per row on body whorl; bases of anteriormost 3 rows of spines joined by low spiral carinae. Posteriormost-but-one (3rd) row of Plate 391. Tectarius rugosus (Wood, 1828). Figs. 1,2. Turbo rugosus Wood, lectotype, BM(NH) 1968370, 28.4 x 22.3 mm. Figs. 3,4. Tectarius coronatus Valenciennes, Holotype, MHNP (“‘Acapulco” [Luzon, Philippines]) 32.9 x 25.7 mim. [05-450] January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Tectarius 515 spines protrudes farthest on spire whorls, but this distinction largely lost on more mature whorls; posteriormost (4th) row of spines obscured by subsutural purplish brown color band especially on penultimate and body whorls. Secondary spiral sculpture, between each row of spines, con- sisting of raised cords, and overall spiral sculp- ture of microscopic threads. Axial sculpture con- sists of irregular flaky lines of growth. Operculum moderate in size, circular, an average one measur- ing about 7 mm. in diameter, thin, light-brown with a dark-brown center, paucispiral, nucleus about central. Periostracum not evident. Nuclear whorls at least partially decollate in all specimens examined, about 2, smooth, grayish white, first postnuclear whorls showing early signs of spiral striae and becoming nodulose. Radula littorinoid, 9-1-1-1-2; lateral tooth with a vertical partition and an embayment typical of Littorinidae. Ani- mal moderately large, littorinoid; penis large, muscular and apparently highly extensible; semi- nal groove in deep fold running along medial edge to tip; tip vermiform, covered with papillae; lateral edge of penis supplied with large number of glands not extending onto vermiform tip. Re- productive data and life history unknown. Measurements (mm) (all Philippines)— length width No. whorls locality 39.7 27.6 7+ Cadao Id., Naro Bay, Masbate 35 25,2 7+ Cadao Id., Naro Bay, Masbate 30.1 22.4 7+ San Miguel Bay, Ticao 95.9 91.2 8 Batag Id., Samar 23.9 16.3 T+ Bongao Channal, SW Sanga Sanga Id., Sulu Archipelago 21.0 15:3. 7+ Borongan, E side Samar io ikea ee Papahag Id., Tawi Tawi Group 15.0 12.0 6+ Borongan Village, E side Samar 13:5 10.8 ate Papahag Id., Tawi Tawi Group 12.6 9.5 6+ Papahag Id., Tawi Tawi La? 8.4 6+ Papahag Id., Tawi Tawi 8.4 6.8 5+ Papahag Id., Tawi Tawi Synonymy— Monodonta papillosa of authors, not M. papillosa Lamarck, 1822 [=Tectarius tectumpersicum (Linne, 1758) ]. 1828 Trochus rugosus Wood, Supplement to the Index Tes- taceologicus or a Catalogue of Shells, British and Foreign, pl. 5, Trochus, fig. 7 (no locality given; Mindanao, Philippines, here selected); lectotvpe in BM(NH) 1968370, length 28.4 mm, width ( ca. ) 22.3; not Litorina rugosa Menke, 1843 [= Nodilittorina aus- tralis (Gray, 1826)]. 1832 Tectarius coronatus Valenciennes in Humboldt and Bonpland, Voyage aux régions equinoxiales du Noveau Continent, vol. 2, Coquilles, p. 271 (Acapulco [in error] locality here corrected to Luzon, Philip- pines); Holotype in MHNP. 1846 Litorina papillosa elegans Philippi, Abbildungen und Beschreibungen Conchylien, vol. 2, p. 140, Litorina, pl. 2, figs. 5, 7 (precise locality not given); figured specimens from Cuming Collection BM(NH) [not seen during 1968 visit] and Saul Collection, Cam- bridge Museum. 1846 Litorina papillosa quadriseriata Philippi, ibid., p. 140, Litorina, pl. 2, fig. 2 (Zanzibar [in error] locality here corrected to Luzon, Philippines); type-specimen may be in BM(NH) [not seen during 1968 visit]; refers to “Trochus rugosus Wood Suppl. t.5, £. 7”. Types—Although the name T. papillosus has been applied to this species (see Kaicher, 1956) the type-specimen of that species in the Geneva Museum is unquestionably T. tectumpersicum Linné, and papillosus is, therefore an absolute synonym of tectumpersicum (q.v.). It also has been referred to as Echinellopsis grandinatus (Habe, 1961, p. 20; 1964, p. 28, both pl. 9, fig. 30) which is an error of nomenclature for T. rugosus. The first available name is Trochus rugosus Wood, 1828, the lectotype of which is in the BM(NH) 1968370. The holotype of T. coronatus Valenciennes is in the Paris Museum. The figured specimens of Philippi’s elegans and quadriseriata may be in the BM(NH) and/or the Cambridge Museum. They were not discovered by me at the BM and may be lost. Philippi’s figures are quite adequate for the interpretation of the species and may be considered as representative of the lecto- types: elegans, pl. 2, fig. 7; quadriseriata, pl. 2, fig. 2. 1oo° 120° \4o° 16¢ . - = ) MARIANA is ANDAMAN , * Soipon Is ‘ * Guar H is niwetok @ Oithi & melo NICOBAR |; cack e 16 % Tt te Pon CEYLON ise, |e RY yiom, Ne LA > COCO6 - KEELING ATOLLS - | | = ee ee ee a pS a te |e een cALE | | = n _L n 100 120° 140° 160 Plate 392. Geographic distribution of Tectarius rugosus (Wood) in the Philippines and Indonesia. [05-451] 516 Joseph Rosewater Records—PHILIPPINES: Port Galera, Mindoro; San Miguel Bay, Ticao Id.; Cadao Id., Naro Bay, Masbate (all USNM); Borongan Village (USNM, Del. Mus. N.H., ANSP, MCZ); Batag Id., both Samar (USNM); Samal Id., Davao Bay (MCZ, USNM, WAM); Zamboanga, both Mindanao (ANSP, Del. Mus. N.H.); Jolo Id. (MCZ); Tabawan Id. (ANSP); Papahag Id. (USNM); Bongao Channel, SW Sanga Sanga Id., all Sulu Archipelago (ANSP). INDONESIA: Bula Buka Id. Gulf of Tomini, Celebes (USNM). Tectarius grandinatus (Gmelin, 1791) (Pl. 388, figs. 1,2; pls. 393-395) Range—The Cook, Society, Tuamotu and Gambier Islands, southeastern Polynesia. Remarks—Tectarius grandinatus, an inhabitant of southeastern Polynesia, apparently is ge- ographically isolated from the several other mem- bers of the genus Tectarius in the Indo-Pacific, the others being found in the East Indies. It is quite distinct, morphologically, from pagodus and tectumpersicum, but, interestingly, is very close in appearance to T. rugosus in general matters of sculpture and external anatomy of the animal. Superficially, grandinatus might be thought more closely related to Cenchritis muri- catus (L.) of the western Atlantic. However, the radula, penial anatomy (Abbott, 1954) and a close examination of shell sculpture and _ structure causes me to reject that theory: grandinatus has not been observed to be umbilicate, while muri- catus sometimes is; the aperture of muricatus is never thickened and plicate as it is in grandinat- us; the operculum of grandinatus is rounded with the nucleus near the center, that of muricatus is more oval with the nucleus near the side. The many similarities between rugosus and grandinatus together with their spatial isolation suggest the possibility that they may have evolved from the same species stock. Although a subspecific relationship may be indicated, the two are here considered to have developed full specific differentiation. Habitat—Usually occurs on low islands within the geographic range; on coral reef flats near the high tide line, on jagged pieces of raised reef. Description—Shell reaching 37.9 mm (about 1 '/l> inches) in length, elongate conical in shape, average obesity about .72 (44 specimens range from .62-.82); rather solidly and thickly con- structed for its size, imperforate; mature speci- mens with very deeply impressed suture and rounded whorls; sculptured with four spiral rows per whorl of stubby spines. External shell color yellowish white; often coated with what appears Littorinidae to be a light-yellow to brown or rather dark grayish brown periostracum which may wear thin especially at tips of spines; no definite color pattern apparent; occasional lines or splotches of medium brown present; upper (most posterior) part of aperture usually covered with a brown glaze. Base moderately flattened, sculptured spi- rally with nodulose cords. Whorls 7-9, moderately rounded. Length of spire usually considerably greater than half the length of shell. Spire convex, produced at an angle of from 58-60°. Aperture rounded; outer lip considerably thickened, pli- cate within, tapering to a thinner, crenulate edge. Inner lip with a thin, brown glaze posteriorly, forming a tooth-like bulge anteriorly, near junc- tion with outer lip near base of columella. Suture often deeply impressed, typically forming a chan- nel between whorls. Primary sculptural feature is the four spiral rows of spines. Spines not regu- larly aligned axially, although anteriormost two rows more-so than others; from about 17-25 spines per row on body whorl; bases of anteri- ormost 3 rows of spines joined by low spiral carinae; bases of posteriormost row of spines usually separate; second from anterior row usual- ly the smallest. Secondary spiral sculpture be- Plate 393. Tectarius grandinatus (Gmelin, 1791). Figs. 1,2. Trochus grandinatus Gmelin, lectotype in ZMC, specimen figured by Chemnitz, Conchylien Cabinet, vol. 10, pl. 169, fig. 1639, from Palmerston Atoll, Cook Is- lands, 32 x 22.4 mm. Figs. 3-5. Monodonta coronaria Lamarck, holotype, MHNG 1096/23, 41 x 27 mm. [05-452] January 15, 1972 tween rows of spines consisting of 2-4 raised cords; overall microscopic sculpture of fine, closely-spaced spiral threads. Axial sculpture consisting of fine, irregular, closely-spaced, over- lapping lines of growth. Shell surface under high magnification may show minute closely-spaced perforations. Operculum moderate in size, rounded-oval, an average one measuring about 9 mm. in diameter, thin, light-brown with a dark- brown center, paucispiral, nucleus about central. A thin, light- to dark-brown periostracum present; closely applied but easily worn away; perios- tracum smooth and dully shining. Nuclear whorls about | !/2, smooth and shining; first post-nuclear whorl rapidly becoming multi-carinate and de- veloping spines on the second. Radula littorinoid, 2-1-1-1-2; lateral tooth with a vertical partition and with an embayment characteristic of Lit- torinidae. Animal moderately large, littorinoid. Penis fairly large and apparently quite extensible; seminal groove in deep fold running along medial edge to tip and bordered by thickened, papillose glandular-appearing tissue; distal end of penis vermiform; lateral edge of penis lined with large number of glands not extending onto vermiform tip. Reproductive data and life history unknown. Measurements (mm)— locality length width no. whorls BYR) 23.4 7+ Manihi, Tuamotu Ids. 34.3 D5 T+ Aitutaki, Cook Ids. 32.0 22.8 6+ Mangaia, Cook Ids. 29.8 9519 7+ Mangareva, Gambier Ids. 94.3 18.7 S+ Bird Id., Palmerston Atoll] 22.0 17.0 9 Bird Id., Palmerston Atoll] 20.7 15.7 7+ Cooks Motu, Palmerston Atoll 18.1 14.7 7+ Mangaia, Cook Ids. 16.1 132 6+ Aitutaki, Cook Ids. 14.3 10.8 6+ Aitutaki, Cook Ids. 12.4 9.1 9 Tikahau Atoll, Tuamotu Ids. 7.8 6.0 6+ Mangaia, Cook Ids. Synonymy— [1784 Trochus bullatus Martyn, The Universal Conchologist, vol. 1, fig. 38; rejected work, I.C.Z.N. Opinion 456]. 1791 Trochus grandinatus Gmelin, Systema Naturae, ed. 13, p. 3585 (ad Palmerstoni insulam [=Palmerston Atoll, Cook Islands); refers to Chemnitz “Conch” vol. 10, p. 291, pl. 169, fig. 1639 and to Martyn “Conch”, vol. 1, fig. 38. Lectotype, here selected, specimen from Spen- gler Collection, ZMC, figured by Chemnitz; see our pl. 393, figs. 1 and 2, 32 x 22.4 mm. 1816 Monodonta coronaria Lamarck, Liste Des Objets Repré- sentés, Tableau Eneyclopédique et Méthodique, part 23, p. 10, pl. 447, fig. 6 a,b (no locality given); Holotype MHNG 1096/23, 41 x 27 mm; 1822, His- toire Naturelle Des Animaux sans Vertébres, vol. 7, Deooe INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 i Tectarius 51 Plate 394. Tectarius grandinatus (Gmelin). Fig. A. Radula. Fig. B. Penis (both from Palmerston Atoll, Cook Islands, USNM 685165). Types—The lectotype of Trochus grandinatus Ginelin, the specimen figured by Chemnitz, vol. 10, p. 291, pl. 169, fig. 1639, is in the Zoological Museum Copenhagen. It measures 32 x 22.4 mm (see pl. 393, figs. 1,2). The Holotype of Monodon- ta coronaria Lamarck is in the MHNG 1096/23 (see pl. 393, figs. 3-5). It measures 41 x 27 mm. 180° 160° 140° | T T T XX \ midway)» Pearl Hermes Haw, Alia Se See ee ts Se eaeteeeceseee a Oohu O3, Woke Pa ‘s ; : | Aces SLHEBRIDES Mois k/ Sate FIJNis Kes NeW bia EDONIA. = oe es . Norfolk | Plate 395. Geographic distribution of Tectarius grandinatus (Gmelin) in the southeastern Pacific Ocean. [05-453] 518 Joseph Rosewater Littorinidae Records—COOK ISLANDS: Palmerston Id.; Aitutaki (both USNM, ANSP, Del. Mus. N.H.); Hervey Ids. (USNM, MCZ, ANSP); E. side Koromiri Id., S.E. Rarotonga (ANSP); Mangaia (USNM). SOCIETY ISLANDS: W. coast Bora Bora (ANSP); Tahiti (USNM, MCZ, ANSP). TUAMOTU ISLANDS: Manihi Atoll (USNM); Takaroa Atoll (ANSP); Maiai Id., Tikehau Atoll (USNM); Aratika Atoll (ANSP); N. of Temao Harbor, Makatea Id. (USNM); Toau Atoll; Raroia Atoll; (all ANSP) Raraka Atoll (Del. Mus. N.H.); Fakarava Atoll; Makemo Atoll; Tatatkoto Atoll; Anaa Atoll; Vahitahi Atoll; Nengonengo Atoll; Tureia Atoll (all USNM); Marutea Atoll, Acteon Group (ZMA). GAMBIER ISLANDS: Mangareva Id. (USNM, MCZ). Tectarius pagodus (Linné, 1758) (Pl. 388, figs. 5-7 Range—From the Philippines, through the Western Pacific Are to the Solomon Islands. Remarks—The name applied by Linné to this species could not have been more appropriately descriptive as the shell with its usually upturned spines very much resembles an oriental turreted pagoda. As pointed out by Dodge (1959) there are a number of similarities between T. pagodus and T. tectumpersicum, both described by Linné. However pagodus reaches a larger size, 21/2 inches versus 1 inch, is proportionately broader and has more prominent spinose carinae than tectumpersicum. There is seldom any difficulty in separating these species with comparative mate- rial at hand. The method of reproduction in pagodus re- mains to be observed. The normal habitat of the species is on cliffs above the sea. As there is no evidence that the species is ovoviviparous, there must be a periodic migration to the sea for breed- ing purposes, although such a phenomenon has not been reported. Another question arising from the high shore habitat of these nominally marine snails concerns their food. Rumphius (1705) early suggested that they feed upon the cliffs where they live. Ex- amination of some fecal pellets of this species revealed the presence of considerable debris and what appeared to be plant cells. It is quite likely that T. pagodus feeds on plant life such as algae and lichens growing on the sea cliffs. The radula is extremely long which may denote such a browsing manner of feeding, the extra length possibly being required because aerial feeding without lubrication from sea water causes a rapid wearing of the teeth (see Quoy and Gaimard, Astrolabe, pl. 62, fig. 1). Habitat—*‘. ... these animals suck their food from the briny moisture of the cliffs to which they cling, being unable to endure the water” (Rumphius, 1705, p. 74). Found on vertical lime- stone cliffs 1-2 meters above high tide line (per- sonal observations, Moluccas Islands, Indonesia, 1970). Description—Shell reaching 61.4 mm. (about 2'/o inches) in length, squatly conical in shape, average obesity about .93 (18 specimens range from .84-1.1); mature individuals rather heavily constructed, imperforate, and sculptured on the body whorl with 2 carinate rows of usually thick, straightly projecting or slightly upturned spines, between which spiral cords are roughly produced and the shell surface often thrown into oblique waves. Overall external color yellowish to grayish white, with no discernable pattern; in young specimens dark-brown spiral lines may be pres- ent externally or within aperture; aperture usually yellowish brown, its edge white. Base distinctly flattened, sculptured spirally with nodulose cords extending into aperture; base separated from upper part of body whorl by one of the rows of spines at periphery. Whorls 5-8, flat-sided, ex- cepting spines. Length of spire usually greater than half the length of shell. Spire convex, pro- duced at an angle, excepting spines, of from 59-65°. Aperture broadly rounded; outer lip thick- ly produced in mature individuals, strongly pli- cate within; plicae not reaching edge of aperture; inner lip smooth, white posteriorly, but forming a prominent tooth-like bulge anteriorly near junc- tion with outer lip at base of columella. Suture somewhat obscure, marked by protrusion of sec- ondary carinae of succeeding whorl. Outstanding sculptural feature is the midwhorl spinose carina, with typically from 9-12 antero-posteriorly flat- tened, triangularly-shaped spines on carina of body whorl. Surface of shell at base of spines raised to form oblique fold often reaching to suture. Secondary spine bearing carinae at per- Plate 396. Tectarius pagodus (Linné, 1758). Fig. 1. Turbo pagodus Linné; lectotype figure from Argen- ville, pl. 11, fig. a. Figs. 2,3. Monodonta bicolor Lamarck, holotype, MHNG 1096/19, 54 x 38.2 mm. [05-454] January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Tectarius 519 iphery of each whorl, entirely visible only on body whorl and protruding above suture of pre- ceeding whorls. Secondary spiral sculpture of raised, roughened spiral cords in turn, covered with countless closely-spaced spiral microscopic textural threads. Operculum large (average about 16 mm. diameter) thin, brown, circular, corneous, paucispiral, nucleus about central. Periostracum not evident. Nuclear whorls partially decollate in all specimens examined, earliest whorl (probably first postnuclear) is spirally striate, the midwhorl carina beginning about second postnuclear whorl; carina becoming nodulose or prespinose almost immediately. Radula littorinoid (2-1-1-1-2) extremely long; lateral tooth partitioned and with an embayment. Animal large, also littorinoid; sides of foot and tentacles bright yellowish orange, remainder of animal grayish brown (color observations on living animals from Kai Islands, Moluccas, Indonesia); penis well-developed, with the seminal duct contained within a deep fold running along its medial edge; with a large number of glands attached along lateral edge; penis minutely papillose also over its surface, unbranched. Reproductive data and life history unknown. Measurements (mm) (width includes spines) locality length width — no. whorls 61.4 45.0 7+ Lutee, Choiseul Id. Solomon Islands 57.1 47.8 6+ Pavuvu Id., Russel Group, Solomon Islands 49.3 42.3 7+ Lutee, Choiseul Id. Solomon Islands 46.9 43.4 8+ Pavuvu Id., Russel Group, Solomon Islands 44.4 42.3 7+ Bougainville Id., Solomon Islands 41.3 37.4 5+ Majugag Id., W coast Buka Id., Solomon Islands 39.7 40.0 8 Lutee, Choiseul Id., Solomon Islands 31.5 31.9 6+ Balagnan Id., Surigao District, Mindanao, Philippines 275 25.5 8 Soepiori Ids., Schouten Ids., West Irian 14.3 12.6 Hf Biak, West Irian Synonymy— 1758 Turbo pagodus Linné, Systema Naturae, ed. 10, p. 762; refers to Argenville, pl. bl, fig. A “Pagodus”’ [lecto- type figure]; (type-locality, O. Asiatico, here restricted to Amboina, Moluccas). 1822 Monodonta bicolor Lamarck, Animaux san Vertébres, vol. 7, p. 31 (no locality); holotype in MHNG 1096/19, 54 x 38.2 mm. 1840 Pagodella major Swainson, A Treatise on Malacology, p. 351 (no locality given); refers to Chemnitz, pl. 163, figs. 1541, 1542 (specimen figured is lectotype, pos- sibly in Copenhagen Museum). 1850 Pagodus verus J. E. Gray in M. E. Gray, Figures of Molluscous Animals, vol. 4, p. 78; refers to Trochus pagodus Quoy [and Gaimard, Astrolabe] pl. [62, not] “82”. figs. 1-4; not Littorina papillosa var. vera Phil- ippi, 1846 [= Tectarius tectumpersicum Linné]. Types—The location of Linné’s type of Turbo pagodus is unknown, and Dance (1967) has pointed out that this species is missing from the Linnaean collection in London. In the absence of a type, one of the figures cited by Linné in connection with the original description is here Plate 397. Tectarius pagodus (Linné). Fig. A. Radula of specimen from West Irian; note smooth cusps probably denoting worn teeth, also “partitioned” lateral. Fig. B. Penis, anterior, and C. posterior views respectively; note glands on lateral edge and papillose surface (both ANSP 207638). | \oo* “ate 5 7S ; 160) J of | > D fl ov 9 > Vy Ohinuvté rise BONIN | L\ 9 = 0 Jim Mare: | oss”) Caaaae ya W renin pela 7 al Saaillpo vce Et | (Oy A a ANDAMAN , ' Soipo “ 1S. ‘ we y *G. | ot 8 NICOBAR | hy L“ CEYLON €0COS - KEELIN Pt ° Sy NEW Qt HEBRIDES ‘ S fo. FILS Ewe i New __ CALEDONIA 9 100° zoe 160° Plate 398. Geographic distribution of Tectarius pagodus (Linné) in the central Indo-Pacific faunal region. [05-455] 520 Joseph Rosewater designated as representing the lectotype: Argen- ville, pl. 11, fig. A. The type-locality, originally given as “QO. Asiatico” is here restricted to Amboi- na, Moluccas. The holotype of Monodonta bicolor Lamarck is in the Museum d’Histoire Naturelle, Geneva (MHNG register no. 1096/19). The loca- tion of types of Swainsons’ Pagodella major and of Gray’s Pagodus verus are unknown to me, and the figures cited in connection with their descrip- tions may be considered as representing their lectotypes (see Synonymy). Records—PHILIPPINES: E. coast Polillo (Del. Mus. N.H., USNM): Calapan, Mindoro Id. (Del. Mus. N.H.); Balagnan Id., Surigao District (USNM); Zamboanga, both Mindanao (ANSP Del. Mus. N.H.). INDONESIA: Pulau We, Sumatra (RNHL); Java (ANSP, RNHL); Timor (RNHL); Morotai Id.; Toetoe Id.; Dagaseli, both N Loloda Group (all MCZ); Ter- nate (RNHL); Buru Id. (ZMA); Ambon (MCZ, ZMA): Tanim- bar Islands (RNHL); Kur Id.; Warbal Id., W of Nuhu Rowa, both Kai Ids. (both USNM, WAM). NEW GUINEA: Waigeo Id. (ANSP); Misool Id.; Fakfak (both Leiden); Manokwari (ANSP); Biak; Soepiori Id., both Schouten Ids. (both USNM); Rouw, Aoeri Ids. (ANSP); Woodlark Id. (MCZ). SOLOMON ISLANDS: Majugag Id., W coast Buka Id.; Nr. Kihili, Buin, Bougainville Id. (both USNM); Choiseul Bay (ANSP); Lutee, both Choiseul Id. (ANSP, USNM): Ataa District, Malaita (ANSP); Roviana (MCZ); Pavuvu, Russell Group (USNM). Tectarius tectumpersicum (Linné, 1758) (Pl. 388, figs. 3,4) Range—From the Philippines along the West- ern Pacific Arc through Melanesia. Remarks—Tectarius tectumpersicum is grossly similar in many ways to T. pagodus and it often appears difficult to construct a point by point comparative description which clearly differen- tiates the two species except in matters of size and degree of obesity; pagodus reaches a length of 21/2 inches (61 mm) while tectumpersicum rarely reaches 1'/2 inches (34 mm); pagodus is very obese, sometimes being wider than high, but in tectumpersicum the width of shell is generally only about 77% of the length. Differences are apparent also in spinosity, there being two rows of spines on the body whorl of pagodus and 3 rows in tectumpersicum. Spines in tectumpersi- cum tend to be stubby and round-ended while in pagodus they are pointed and broadly triangular, although there are occasional specimens of tec- tumpersicum which tend to resemble small, ma- ture pagodus. Generally, however, specimens of pagodus the size of tectumpersicum are obvious- ly immature and thin-lipped, so that the “rule-of- thumb” involving size of specimens can be de- pended on to separate the species. Littorinidae Habitat—Lives in pockets of worn, raised lime- stone reef rock, 1-2 meters above high tide line (personal observations, Moluccas Islands, In- donesia, 1970). Description—Shell reaching 34.6 mm (about 1*/16 inches) in length, conical in shape, average obesity about .77 (38 specimens range from .64 ~.85), mature individuals moderately thick in structure, imperforate, and sculptured with three main rows of stubby, often upturned, spines on body whorl, and usually with two rows on spire whorls, between which spiral cords are roughly produced, wavy or papillose and often approach- ing minor rows of spines. External color yellow- ish to grayish white, with no regular patterning although some specimens have diffuse dark spiral color bands externally or within aperture es- pecially at edge of outer lip or on tooth-like bulge of inner lip. Remainder of aperture usually white or yellowish white. Base moderately flattened, Plate. 399. Tectarius tectumpersicum (Linné, 1758). Figs. 1-3. Turbo tectumpersicum Linné, lectotype in Lin- nean Society of London collection, ca. 25 x 23 mm. Figs. 4-7. Monodonta papillosa Lamarck, lectotype, MHNG 1096/22-2, 29.3 x 25.5 mm. [05-456] January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Tectarius 52] Plate 400. Tectarius tectumpersicum (Linne). Fig. A. Radula. Fig. B. Penis (both from Biak, West Irian, ANSP 206421 and USNM 637390). sculptured spirally with nodulose cords which may extend into aperture; base separated from upper part of body whorl by a row of spines at periphery. Whorls 6-8, rather flatsided excepting spines. Length of spire greater than half the length of shell. Spire convex, produced at an angle of from about 55-65°. Aperture compactly rounded; outer lip thickly produced; strongly plicate within, but thin and crenulate at apertural edge; inner lip smooth posteriorly, but forming a tooth-like bulge anteriorly near junction with outer lip at base of columella. Suture obscure, partially masked by lower row of spines at pe- riphery of whorls. Center row usually bearing largest spines, from 9-12 on body whorl; the more posterior row, near suture has smaller spines but about same number; anterior row at periphery smaller still and more numerous with from 13-16 spines. On spire whorls center row of spines may predominate with others being either hidden or suppressed. Spines usually not aligned axially. Secondary spiral cords in some specimens almost as spinose as primary ones; in other specimens hardly noticeable. Entire surface covered with closely spaced spiral microscopic — textural threads. Axial sculpture consists of often coarse, irregular lines of growth. Operculum moderate in size (average about 6-7 mm diameter), thin, brown, circular, corneous, paucispiral, nucleus about central. Periostracum not evident. Nuclear whorls about two, but at least partially decollate in all specimens examined; brown, smooth for at least 1 volution, then becoming carinate; first post nuclear whorl weakly nodulose and rapidly be- coming spinose. Radula littorinoid (2-1-1-1-2) [teeth about !/2 the size of those of T. pagodus]; lateral tooth partitioned and with an embayment. Animal medium-sized, littorinoid; penis large, unbranched, with a large number of glands along lateral edge, 3/4 the length to tip; surface of penis otherwise papillose; seminal duct deeply folded. Life history unknown. Measurements (mm) (width includes spines)— locality length width no. whorls 34.6 24.5 6+ Pavuvu Id., Russell Group, Solomon Islands 31.4 Bo 8 Lunga, Guadalcanal, Solomon Islands 96.7 14 7+ Timor, Indonesia 22.8 18.4 6+ Cebu, Philippines 21.4 15.9 7+ Biak, West Irian 20.0 5:5 6+ Biak, West Irian 19.2 16.0 6+ Cebu, Philippines 18.1 13:7 7+ Biak, West Irian 15.2 12.9 6+ Anir Id., New Ireland 13.6 10.3 8 Philippines Synonymy— 1758 Turbo tectumpersicum Linné, Systema Naturae, ed. 10, p. 762 (no locality given; Cebu Id., Philippines, here selected as type-locality); lectotype in Linnean Soci- ety of London collection. —— Trochus bullatus “Martvn” of authors; [not T. bullatus Martyn, 1784, Universal Conchologist, vol. 1, fig. 38; non-binomial; is Tectarius grandinatus (Gmelin, 1791)]. 1822 Monodonta papillosa Lamarck, Histoire Naturelle des Animaux sans Vertébres, vol. 7, p. 32. (“les mers de Timor’); lectotype MHNG 1096/22-2. 1846 Litorina papillosa vera Philippi, Abbildungen und Be- schreibungen Conchylien, vol. 2, p. 141 (no locality given); refers to “Delessert, Recueil, pl. 36, fig. 10,” here selected as the lectotype figure. 100° 120° (40° _160 180 “MARIANA, Wome ohnston ewe, TECTUMPERSICUM | «2° "MARSHALL IS 2 = 7 = oo © y . = TA ve ies — 'y @ 5 ee y ; PHOENIX 18 “Wey TOKELAU 1S Sys *. A S : ; 3 ” Aton ts > 3 new > x y HEBRIDES : et 3 = |e by, FIIs NEW TONGA CooK a CALEDONIA cd 1s a 1 T | Nor fe i] NEW | ZEALAND | Le | | n ale 1 | 1 & 10° 120° 140° 160° Tao" Plate 401. Geographic distribution of Tectarius tectumper- sicum (Linne) in the central Indo-Pacific faunal region. [05-457] O22 Joseph Rosewater Littorinidae Types—The specimen of Turbo tectumper- in structure, imperforate, and sculptured with 2-3 sicum Linné in the Linnaean shell collection located in the Linnean Society, London, may be considered the lectotype of this species; it mea- sures approximately 25mm length (about | inch). The lectotype somewhat resembles T. pagodus but may be recognized by its rounded spines and the mature, thickened shell. The figure cited by Linné, “Argenville, pl. 11, fig. P’’, is controversial and a poor representation of the species (also see Dodge, 1959, p. 229). As no type-locality was given by Linné, Cebu Island, Philippines, is here selected as a place from which specimens could have come in the early 18th century. The figure referred to by Philippi, “Delessert, Recueil, pl. 36, fig. 10”, is here selected as the lectotype figure of Litorina papillosa vera. The lectotype of Mon- odonta papillosa Lamarck, which proves to be an absolute synonym of T. tectumpersicum is in the Geneva Museum (MHNG 1096/22-2; see our pl. 399 figs. 4-7). Nomenclature—The tectumpersicum does not need to agree in gender with Tectarius because the former is a noun in apposition, mean- name ing “persian roof”. Records—PHILIPPINES: Cebu (USNM). INDONESIA: Timor (USNM, RNHL); Ambon; Kur Id.; Warbal Id., W of Nuhu Rowa, both Kai Islands (all USNM, WAM). NEW GUINEA: Wasior, Wandammen Bay (ZMA); Soepiori Id., (MCZ): Biak Id., both Schouten Islands, all West Irian (USNM); reef at Cape Moem, nr. Wewak, Territory of New Guinea (MCZ). BISMARCK ARCHIPELAGO: Feni Ids., E of New Ireland (USNM). SOLOMON ISLANDS: Nissan Id., Green Islands (SMF); Choiseul Bay, Choiseul Id. (ANSP); Stirling Isle, Treasury Ids.; Munda, New Georgia; Pavuvu Id., Russell Group; Lunga, Guadalcanal (all USNM); Santa Ana Id., S. of San Cristobal (ANSP). Tectarius rusticus (Philippi, 1846) (Pl. 388, figs. 8,9; pls. 402, 403) Range—Northern Western Australia. Remarks—This species is restricted to northern Western Australia according to available locality data. It appears to be most closely related to T. rugosus trom which it differs in its usually less regularly sculptured, more flat-sided whorls and in its lack of external coloration. In T. rusticus the subsutural, midwhorl and peripheral rows of spines are quite commonly strongly expressed, while in rugosus all rows are subequal. Habitat—Lives on rocks above high tide line. Description—Shell reaching about 40 mm (about 1°/s inches) in length, broadly conical in shape, average obesity about .77 (29 specimens range from .72-.83); becoming moderately thick main rows per whorl of rather muted spines. External color yellowish white with occasional faint orange stripes; inside of aperture white and occasionally tinged with pinkish orange, often with brown lines at its edge and revolving within; apertural tooth often stained with brown. Base flattened, sculptured spirally with nodulose cords, the strongest of which occurs just below periphery of body whorl. Whorls 6-8, rather flat sided. Length of spire greater than half the length Plate 402. Tectarius rusticus (Philippi, 1846). Figs. 1,2. Litorina papillosa rustica Philippi, knobby and smooth forms, respectively. Fig. 2 (lectotype figure) probably is representation of Phi- lippi’s concept of L. p. subinermis, the smooth form; both from Point Swan, northern Western Australia; from Ab- bildungen und Beschreibungen Conchylien, vol. 2, Litorina, pl. 2, figs. 3,4. [05-458] January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Tectarius 523 of shell or the two may be about subequal. Spire convex, produced at an angle of about 68°. Aper- ture rounded to squarish; outer lip thickly pro- duced, internal plicae only moderately produced and not reaching edge of aperture; outer lip tapering to thin crenulate edge; inner lip forming a tooth-like bulge anteriorly near junction with outer lip; tooth often stained with brown. Suture usually obscured by anteriormost row of spines of preceding whorl. Spiral spinose sculpture sub- dued, usually three rows are outstanding: the anteriormost, posteriormost and central rows of each whorl, the rest being limited to undulating, slightly bumpy cords; from 12-14 spines per row on body whorl; spines sometimes coalesce into oblique axial bars. Fine sculpture composed of microscopic wavy spiral cords and finer threads. Operculum moderate in size, circular, light- brown with a dark-brown center, paucispiral, nucleus about central. Periostracum not evident. Nuclear whorls partly decollate in all specimens examined; remaining portions smooth, white; early postnuclear whorls rapidly becoming spi- rally striate. Radula littorinoid, 2-1-1-1-2, similar in appearance to that of T. rugosus. Preserved specimens not available for observations on anat- omy; radula obtained from dried specimen. No- thing reported concerning reproduction and de- velopment. Measurements (mm) (all Western Australia)— length width no. whorls locality 35.5 29.0 5+ Yampi Sound, W.A. 34.5 D7A5 6+ Cliff Id., King Sound 32.4 2319 8+ Buccaneer Archipelago 25.8 20.6 7+ Buccaneer Archipelago D322; 18.8 5+ Cliff Id., King Sound 22.1 17.1 6+ Buccaneer Archipelago 21.9 16.2 7+ Buccaneer Archipelago 21.1 15.3 7+ Cliff Id., King Sound 16.3 12h 6+ Buccaneer Archipelago 15.8 1:22, 6+ Buccaneer Archipelago Synonymy— 1846 Litorina papillosa rustica Philippi, Byeuiluaen und Beschreibungen Conchylien, vol. 2, Litorina, p. 140, pl. 2, fig. 3 [and 4 Lectotype Fe (Point Swan [Cape Leveque, Dampier Land, northern Western Australia]; type-specimen may be in Berlin Museum). 1846 Litorina papillosa subinermis Philippi, ibid., p. 141. pl. 2, fig. 4 1857 Littorina bullata in Reeve, Conchologia Iconica, vol. 10, Littorina, pl. 1, fig. lc; not Trochus bullatus Martyn [non-binomial] which is Tectarius grandinatus Gmelin. 1971 Tectarius pagodus Linnaeus, Wilson and Gillett, Aus- tralian Shells, p. 30, pl. 11, fig. 9. Records—WESTERN AUSTRALIA: Troughton Islands, N. a Admiralty Gulf; S.E. Wood Island, E of Cockatoo Island: Yampi Sound; Koolan Island (all WAM); Kellan Island, Buc- caneer Archipelago (AMS; USNM; MCZ): Cliff Island (USNM) all vicinity of King Sound. + iz MARIANA > 1s f oe | psy, PHILIPPINE * Guana | Brio . % ig te . "Ss, Yop , 1 Uhthi Envwetok Polou 4 * : or pines re 2 a elena eA AI * +» Ponape CAROLINE “hes f° 100° yzo0° 140° 160° Plate 403. Geographic distribution of Tectarius rusticus (Philippi) in northern Western Australia. Tectarius songoense (Martin, 1931) (Pl. 404, figs. 3,4) Range—Eocene of Java, Nanggoelan-beds, Kali Songo. Remarks—It is not too difficult to determine from Martin’s illustration just why he assigned this species to “Tectarium”. There is a weakly nodulose sculpture, perhaps better described as beaded, and there appears to be a columellar tooth, a structure characteristic of Tectariinae. The general shape and appearance are, however, more suggestive of Trochidae. The species is tentatively here placed in the subfamily Tectari- inae for want of positive proof to the contrary. Synonymy— 1931 Tectarium (Echinella) songoense kK. Martin, Wetens- schappelijke Mededeelingen Dienst Mijnbouw, no. 18, p. 41, pl. 6, fig. 5, 5a, (Upper Eocene, Nanggoe- lanbeds, Java); unique holotype probably in Geologi- cal Museum, Leiden; length $8 mm. [05-459] 524 Joseph Rosewater Littorinidae Subgenus Subditotectarius Ladd, 1966 Type: Tectarius rehderi Ladd, 1966 Small, stout; spire conical, base convex; with beaded spiral ribs and slightly oblique axial lines; aperture strongly plicate within. Sculpture sub- dued, consisting of small beads formed by con- vergence of axial growth lines and spiral threads. Synonymy— 1966 Subditotectarius Ladd, Geological Survey Professional Paper 531, p. 59; type-species by original designation: Tectarius rehderi Ladd. Tectarius (Subditotectarius) rehderi Ladd, 1966 (Pl. 404, figs. 5-7) Range—Lower Miocene, Marshall Islands. Remarks—The fossil Subditotectarius rehderi is characterized by its subdued beaded sculpture as compared with Tectarius s.s. with its larger nodules and spines. Otherwise the species is quite similar to other Tectarius in outline and sculpture, and in the presence of a columellar tooth and plications within the aperture. Al- though apparently clearly a tectariine S. rehderiis not believed to be closely related to any living species. Synonymy— 1966 Tectarius (Subditotectarius) rehderi Ladd, Geological Survey Professional Paper 531, p. 59, pl. 11, figs. 11-13 (drill hole 2A, Bikini Atoll, 1,051-1,057 feet; early miocene); holotvpe USNM 648342, 2.8x 2.4mm. Plate 404. Fossils of Tectariinae and Echinininae. Figs. 1,2. Echininus adelaidensis (Cotton, 1947) holotype, 16 x 12 mm. Figs. 3,4. Tectarius songoense Martin, 1931; holotype, 8 mm length. Figs. 5-7. Tectarius rehderi Ladd, 1966, holotype, 2.8 « 2.4 mim. [05-460] January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Subfamily Echinininae, new subfamily Genus Echininus Clench and Abbott, 1942 Type: Echininus cumingi (Philippi, 1846) The genus Echininus sensu lato includes the nominate subgenus, Echininus, typified by the umbilicate and spinose EF. cumingi and also E. cumingi spinulosus, both of the western Pacific, and the subgenus Tectininus Clench and Abbott, 1942, having the non-umbilicate and less spinose monotype, E. (T.) nodulosus (Pfeiffer), a species which inhabits the Caribbean area. Members of both subgenera have what may be called multi- spiral (polygyrous spiral type) opercula and pos- sess moderately spinose shells. In Echininus sensu stricto the central radula tooth is moder- ately reduced in width, while in Tectininus it is dramatically reduced and narrowed. Abbott (1954) discussed the phylogenetic position of Echininus and concluded that it belongs in Lit- torinidae although it possesses many specialized characters such as the multispiral operculum, an umbilicus and a narrow central radula tooth. As there are no representatives of Tectininus in the Indo-Pacific, only Echininus s.s. will be con- sidered here. Synonymy— 1854 Echinella ‘Swainson’ H. and A. Adams, The Genera of Recent Mollusca, vol. 1, p. 316; three species men- tioned: coronaria Lamarck [= Tectarius grandinatus Gmelin]; granulata Swainson [unrecognizable]; and the exemplary species, cumingii Philippi; 1895, Pils- bry, Catalogue of the Marine Mollusks of Japan, p. 175; 1901, Pilsbry, Proceedings of the Academy of Natural Sciences of Philadelphia, p. 198; 1903, Kes- teven, Proceedings of the Linnean Society of New South Wales, for 1902, part 4, p. 632, in part; not Echinella Swainson, 1840 [= Tectarius s.s.]. 1942 Echininus Clench and Abbott, Johnsonia, vol. 1, no. 4, p. 3; new name for Nina Gray, 1850, Figures of Molluscous Animals, London, vol. 4, p. 78; Type species by monotypy Trochus cumingii Philippi; not Nina Horsfield, 1829, nor Gray, 1855. Subgenus Echininus s.s. Pyramidal, umbilicate littorinids with a stron- gly spinose shell. In males the penis has a deep but open sperm duct, and basal penial glands are present. Operculum multispiral. Radula littori- noid, the central tooth moderately narrow. Echininus 525 Echininus cumingi cumingi (Philippi, 1846) (Pl. 388, figs. 12,13) Range—From the Philippines along the west- ern Pacific arc to New Hebrides and the Cook Islands. Remarks—At first examination one finds it sur- prising that “Cuming’s Echininus” is included in the Littorinidae. This species has many of the attributes of certain other families, perhaps re- sembling most some members of the family Tro- chidae with its conical shape, multispiral opercu- lum and well-defined umbilicus. Animal charac- ters, especially the radula, and gross reproductive features leave no doubt, however, that E. cumingi is a littorinid. Its closest relative is the smaller and less spinose, but otherwise very similar ap- pearing subspecies, EF. cumingi spinulosus (Phil- ippi). The next most closely related and only other species in the subfamily, the Western Atlan- tic E. (Tectininus) nodulosus (Pfeiffer) differs in never being umbilicate and in having a consider- ably more reduced radula. The open condition of the spines occurs occasionally in E. nodulosus and their arrangement or alignment is similar. Both have multispiral opercula. Penial anatomy differs, however, E. cumingi having a cluster of basal penial glands, whereas E. nodulosus dis- plays a basal swelling and a single gland located one half to two thirds the distance to the tip (see Abbott, 1954, fig. 55, p,q). Habitat—Lives in pockets of weathered, raised limestone reef, 3-7 meters above high tide line (personal observations in Davao, Philippines and Moluccas Islands, 1970). Description—Shell reaching 20.4 mm (about 3/4 inch) in length, broadly conical in shape, with Plate 405. Echininus cumingi cumingi (Philippi, 1846). Trochus cumingii Philippi, lectotype figure of specimen in BM(NH), from Reeve, 1857, Conchologia Iconica, vol. 10, Littorina, pl. 2, fig. 8. [05-467] 526 Joseph Rosewater Littorinidae projecting spines, average obesity about .88 (33 specimens range from .77 to 1.03) only moder- ately thick in structure, umbilicate; suture im- pressed, whorls slightly rounded; sculptured with three rows of short, projecting, often un- closed spines. External shell color grayish tan, the spines often being reddish brown; a thin grayish brown periostrical coating apparent; aperture a diffuse yellowish to reddish brown, occasionally with three reddish brown color bands reflecting the position of the external spines. Base distinctly flattened, spirally sculptured with nodulose cords of which about the third below the periphery is the strongest. Umbilicus very deep but narrow in mature specimens; bordered medially by a rim of the columellar callus and distally by a basal fold. Whorls 6-8, only slightly rounded. Length of spire greater than half the length of the shell. Spire convex, produced at an angle of from about 66-76°. Aperture rounded, outer lip only moder- ately thickened, smooth within, the edge often undulating in the vicinity of the rows of spines; inner lip curved and shining, edentulous. Suture usually somewhat obscured by anteriormost row of spines of preceding whorl. Primary sculptural feature is the three spiral rows of spines. Posteri- ormost row of spines consisting of low, rounded protuberances; anteriormost row small, unclosed, moderately projecting, narrow-hoodlike and closely spaced; middle row of spines largest, most projecting, also incompletely closed. Rows. of spines not aligned axially; spine count as follows: anterior row, 18-25; middle row, 14-17; posterior 15-16 (approximate range). Secondary spiral sculpture between rows of spines consists of from 4-6 slightly raised cords; overall microscopical spiral sculpture of fine, closely spaced threads. Axial sculpture consists of fine irregular often overlapping lines of growth. Operculum moder- ate in size, multispiral (polygyrous spiral type) having about 5-7 volutions, chitinous, dark- brown (pl. 389). Nuclear whorls smooth, light brown, shining, about 1.5 volutions; postnuclear whorls rapidly becoming striately sculptured and then spinose. Radula littorinoid, 2-1-1-1-2, central tooth very narrow, rather simple; lateral tooth only moderately narrow and with a well devel- oped littorinoid notch. Animal littorinoid; penis large and_ well- developed, with a bulbous swelling at its base; distal extremity simple; with as many as 12 penial glands clumped mostly on posterior surface near junction of bulbous base and extremity; sperm Plate 406. Echininus cumingi (Philippi). Fig. A. Radula; note “spurs” on central tooth, and generally narrow dentition. Fig. B. Penis, 1. anterior, 2. posterior, 3. cross-section of seminal groove; note cluster of glands near bulbous base (both from Davao Gulf, Mindanao, Philippines). groove deeply folded, with an apparent internal fold best seen distally. Details of reproduction and life history unknown. Measurements (mm)— length width no. whorls locality 20.4 Lio 7 Cook Islands 19.5 17.6 8 Philippines 18.5 14.9 7+ Stirling Isle, Treasury Ids., Solomons 17.5 14.3 7+ Stirling Isle, Treasury Ids., Solomons 17.0 14.9 Cook Islands 16.7 15.3 7+ Cook Islands 16.3 14.8 7+ Stirling Isle, Treasury Ids. Solomon Ids. 15.9 12.3 7+ Stirling Isle, Treasury Ids. Solomon Ids. 15.1 13.0 7+ Tana, New Hebrides 14.6 13.7 6+ Stirling Isle, Treasury Ids. Solomon Ids. 13.2 11.9 7+ Stirling Isle, Treasury Ids. Solomon Ids. 10.6 10.9 8 Philippines Synonymy— 1846 Trochus cumingii Philippi, Proceedings of the Zoologi- cal Society of London, for 1845, p. 138 (Guimaras Id. {south of Panay Id.] Philippines; as Litorina, 1847, Abbildungen und Beschreibungen Conchylien, vol. 3, [05-468] January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Echininus 527 Litorina, p. 53, pl. 6, fig. 22; lectotype in BM (NH), figured by Reeve, 1857, Conchologia Iconica, vol. 10 Littorina, pl. 2, fig. 8. 1879 Trochus echinulatus ‘Kiener’ in P. Fischer, Spécies Général et Iconographie des Coquilles Vivantes, pl. 43, figs. 2 [name and figure only]; ibid., p. 459, places name in synonymy of Tectarius cumingi (sic); not Trochus echinulatus A. Alth, 1850. Records—PHILIPPINES: Samal Id., Davao Bay, Mindanao (USNM, MCZ). INDONESIA: Karakelong Id., Talaud Ids. (MCZ); N. shore Warbal Id., W of Nuhu Rowa, Kai Ids; W. side Mitak Id., Jamdena Strait, Tanimbar, both Moluccas (USNM, WAM). NEW GUINEA: Misool; Fakfak (both RNHL,); Louisade Ids. (RNHL, NMW, ANSP). SOLOMONS: Stirling Isle, Treasury Ids., (USNM); Santa Ana (ANSP). NEW HEBRIDES: S end Black Beach, Tana (USNM). COOK ISLANDS: Mauke, Hervey Ids.; Rarotonga (both ANSP); Mangaia (USNM). TUAMor, Plate 407. Geographic distribution of Echininus cumingi (Philippi) in the East Indies and Pacific Ocean, and of its more northerly distributed subspecies E. cumingi spinulosus (Philippi). Echininus cumingi spinulosus (Philippi, 1847) (PI. 388, figs. 14,15; pls. 407, 408) Range—From southern Japan through the Ryu- kyu Islands, the northern and western Philip- pines and eastward to the Mariana Islands. Remarks—Echininus spinulosus is very close in its relationship to E. cumingi, the differences between the two being more of degree than of kind. The two are apparently geographically iso- lated, or at least they occupy separate ranges and so the phenotypic differences may be ecologically influenced as well as having a genetic basis. Whatever the basis for the differences between them, it seems appropriate to consider them as subspecies. Echininus spinulosus never reaches as large a size or achieves the squatly conical shape of E. cumingi, and although the sculpture of the two is basically very similar, cumingi is always more distinctly spinose. Both are usually umbilicate although spinulosus is often narrowly so and young specimens may lack this feature entirely, as did Philippi’s type-specimen. It is interesting to note that E. luchuana was described by Pilsbry (1901) as a subspecies of cumingi, which has caused some confusion. Some malacologists have continued to use the combination E. cumingi luchuana or simply E. cumingi when referring to the entity E. spinulo- sus (Kira, 1959, 1962; Habe, 1951). Philippi’s figures (1847) clearly show the species concepts he intended: E. spinulosus (fig. 24), the smaller less pronouncedly spinose species (which in- cludes luchuana as a synonym) and EF. cumingi (fig. 22), the larger and more outstandingly spi- nose and more squatly conical species. Habitat—On rocks above the high tide line. Description—Shell reaching 16.8 mm (about 5/s) inch in length, turbinate in shape, with short spines; average obesity about .84 (32 specimens range from .74-1.08) moderately thick in struc- ture, mature specimens usually umbilicate; su- ture evident although often obscure; whorls mod- erately rounded; sculptured with three major rows of short spines per whorl. External shell color grayish to tannish orange, the short spines usually appearing whitish; apertural coloration Plate 408. Echininus cumingi spinulosus (Philippi, 1847) Figs. 1-3. Echinella cumingi luchuana Pilsbry, lectotype, ANSP 70962, 16 x 14 mm. Fig. 4. Litorina spinulosa Philippi, lectotype figure, from Abbildungen und Beschreibungen Conchylien, vol. 3, Litorina, pl. 6, fig. 24. [05-469] 528 Joseph Rosewater Littorinidae tannish to brownish orange, occasionally with 3-4 darker brown color bands revolving within. Base moderately flattened, sculptured with nodulose cords of which about the third below the per- iphery is the strongest. Umbilicus usually rather narrow and occasionally absent, bordered medi- ally by a rim of the columella callus and distally by a poorly to well defined basal fold. Whorls 6-8, only moderately well rounded. Length of spire greater than half the length of the shell. Spire convex, produced at an angle of from about 63-68°. Aperture rounded-oval, outer lip thin to only moderately thickened, smooth within, and often slightly undulating in vicinity of external spine rows; inner lip curved and shining, edentu- lous. Suture fairly evident to somewhat obscured by anteriormost row of spines of preceding whorl. Primary sculptural feature is three spiral rows of spines located centrally, anteriorly and posteri- orly on each whorl; spiral cords between main rows tending to become nodular to spinose. Rows of spines not well aligned axially; spine count as follows: anterior row about 24; middle row about 19-23; posterior row about 16-19. Secondary sculpture consisting of 4-5 rows of spiral cords which occasionally become nearly as strongly spinose as the 3 primary rows which they sepa- rate; overall microscopical spiral sculpture of fine, closely-spaced threads. Axial sculpture con- sists of fine irregular lines of growth. Operculum small to moderate in size, multispiral (polygyrous spiral type) having 5-6 volutions, chitinous, dark brown. Nuclear whorls smooth, tannish white, shining, about 1.5 volutions; postnuclear whorls rapidly becoming striated and nodulose. Radula littorinoid, 2-1-1-1-2; central tooth narrow and reduced; lateral tooth with a distinct littorinoid notch. Animal littorinoid; penis unbranched; a well developed sperm groove running along its medial edge; distal portion papillose; 2-3 penial glands located half-way between base and tip in preserved specimen. Nothing has been reported concerning reproduction and life history. Measurements (mm)— length width no. whorls locality 16.8 11.7 8 Fuga Id., Philippines 15.6 12.2 7+ “Japan” 14.9 Val T+ Yokohama, Japan 14.2 11.8 7 Batan Id., Philippines 13.9 11.5 6+ Batan Id., Philippines 13.5 10.9 8 Batan Id., Philippines 13.0 10.4 7 Kume-shima, Ryukyu Ids. 12.6 10.4 6+ Okinawa, Ryukyu Ids. 12.0 10.0 7+ Batan Id., Philippines 11.1 9.0 a Batan Id., Philippines 10.8 10.5 7 Batan Id., Philippines 7.3 7.9 6 Batan Id. Philippines Synonymy— 1847 Litorina spinulosa Philippi, Abbildungen und Beschrei- bungen Conchylien, vol. 3, Litorina, p. 53, pl. 6, fig. 24, Lectotype Figure (Manila); type may be in Berlin Museum. 1895 ‘Echinella cumingi Phil. in Pilsbry, Catalogue of the Marine Mollusks of Japan, published by Frederick Stearns, Detroit, p. 175 (Yaeyama [Okinawa)]). 1901 Echinella cumingi luchuana Pilsbry, Proceedings of the Academy of Natural Sciences of Philadelphia, p. 198 (Loo Choo Islands [Ryukyu Islands]); ibid. p. 394, pl. 19, fig. 16; lectotvpe ANSP 70962, ca. 16 « 14 mm. Records—JAPAN: Yokohama (USNM). RYUKYU_ IS- LANDS: Kadena Circle (USNM):; Tsukin-shima, both Okina- wa (MCZ): Kume-shima (MCZ ANSP, USNM, BPBM); Kari- mata, Miyako-shima (MCZ, ANSP); Ora Wan (USNM). TAI- wan; Hung-t?ou Hsu, off Pacific Coast (ANSP, USNM). PHILIPPINES: Santa Domingo de Basco, Batan, Batan Ids. (USNM); Dalupiri Id. (MCZ); Fuga Id., both Babuyan Ids. (USNM); Camp Wallace, Province of La Union, Luzon, Puerto Princessa, Palawan (both USNM). MARIANA IS- LANDS: Saipan; Piti Bay, Guam (both ANSP); Apra Bay; Asan Point, both Guam (both USNM). Echininus adelaidensis (Cotton, 1947) (Pl. 404, figs. 1,2) Range—Adelaidean (Pliocene), South Aus- tralia. Remarks— Echininus adelaidensis certainly is a unique appearing species which somewhat re- sembles certain of the Trochidae nearly as much as it does Echininus (cf. Turcica A. Adams or Perrinia H. and A. Adams, as shown in Wenz, 1938). The characteristics of its partly open spines and the presence of an umbilicus may be sufficient to relate it to Echininus, however. Cot- ton refers to E. cumingi Philippi having been collected in Caloundra, Queensland and Western Australia, but I have not seen such records in the course of the present study. It is possible that northern portions of Australia may be within the range of E. cumingi which reaches the southern Moluccas and New Guinea. Synonymy— 1947 Nina adelaidensis Cotton, Records of the South Austra- lian Museum, vol. 8, no. 4, p. 666, pl. 21, figs. 17, 18 (Adelaidean Pliocene, Salisbury Bore, 350 feet); holo- type in Tate Museum, University of Adelaide, 16 « 12 mm. [05-470] INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 January 15, 1972 [replacing vol. 2, no. 11, pp. 453, 454; see stars] Littorina 529 “Littorina” incisa Yokoyama, 1927 (Pl. 358, figs. 4, 5) Range—Pliocene of Japan. Remarks—As pointed out by Habe (in litt., 1971), this 5 mm. shell is a member of the Pyramidellidae. We erroneously considered it to be a Littorina (Littorinopsis) in our last number of Indo-Pacific Mollusca, vol. 2, no. 11, p. 466 [p. 052340]. pl 358, fase4, Dp. Synonymy— 1927 Littorina incisa Yokoyama, Journal of the Faculty of Science, Imperial University of Tokyo, section I, Geology, Mineralogy, Geography, Seismology, vol. 2, part 4, p. 175, pl. 47, fig. 8 (Pliocene, Nagaya, Kaga, Japan); holotype in Geological Institute, Imperial University of Tokyo: 5 « 2.5 mm. 1970 Littorina incisa Yokoyama, Rosewater, Indo-Pacific Mollusca, vol. 2, no. 11, p. 466. Littorina kozaiensis Nomura and Onisi, 1940 (Pl. 349, figs. 6, 7) Range—Lower Miocene of Japan. Remarks—This species was described as re- sembling L. adonis Yokoyama (see below), but as having a larger number of spiral grooves. The unique holotype (pl. 349, figs. 6, 7, a copy of the original illustration) offers little basis for com- parison with Recent species. Synonymy— 1940 Littorina kozaiensis Nomura and Onisi, Japanese Jour- nal of Geology and Geography, vol. 17, nos. 3 and 4, p. 191, pl. 19, fig 6 a,b. (Y6suibori, Simizu, Kozaimura, Japan); holotype: Saito H6-on Kai Museum, Register No. 21762; 11 x 8 mm. Littorina adonis Yokoyama, 1927 (Pl. 349, figs. 8, 9) Range—Pliocene of Japan. Remarks—Placed provisionally here in the subgenus Littoraria, this species resembles L. undulata, although the strong spiral sculpture is also reminiscent of Littorinopsis, i.e. L. scabra, etc. Unfortunately the outer lip and a portion of the body whorl of the type (pl. 349, figs. 8, 9) are missing and it is difficult to be sure of their exact shape. Synonymy— 1927 Littorina adonis Yokoyama, Journal of the Faculty of Science Imperial University of Tokyo, section 2, vol. 1, part 10, p. 451, pl. 51, fig. 8, (Upper Musashino, Koyasu southern Musashi, Japan); (unique holotype in collection of Geological Institute Imperial Univer- sity of Tokyo: 6 x 4 mm.). Littorina lucida Yokoyama, 1927 (Pl. 349, figs. 4, 5) Range—Pliocene of Japan. Remarks—This species is from the same de- posit as L. adonis but lacks the deeply incised spiral sculpture. The type of lucida, although of approximately the same size as adonis is more slender. We overlooked the fact that Habe, 1942 (Venus, vol. 12, p. 37) and Abbott, 1958 (Proc. Acad. Nat. Sci. Phila., vol. 110, p. 270) had pointed out that L. lucida was a synonym of Assiminea japonica von Martens, 1877. Synonymy— 1927 Littorina lucida Yokoyama, Journal of the Faculty of Science Imperial University of Tokyo, section 2, vol. 1, part 10, p. 451, pl. 51, fig. 9 (Upper Musashino, Koyasu southern Musashi, Japan; unique holotype in collection of Geological Institute Imperial University of Tokyo: 5 * 3 mm.). [05-317a] 530 Littoraria Joseph Rosewater Littorinidae [ These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.] [05-318a] January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Index 531 INDEX TO LITTORINIDAE NAMES IN VOL. 2, NOS. 11 and 12 Looseleaf subscribers should place this index at the begin- ning of the family Littorinidae, and just after the index tab Littorinidae, p. 05-250. The number following the name refers to the pagination found at the top of the page. The column at the right is the looseleaf pagination. All new names are in boldface type. [looseleaf] acuminata Gould, 438 05-300 acuta Menke, 469 05-353 acutispira E.A. Smith, 451 05-312 adelaidensis Cotton, 528 05-470 adonis Yokoyama, 453 05-317 afer Férussac, 431 05-279 africana Philippi, 476 05-360 albicans Metcalfe, 455 05-329 459 05-333 Algaroda Dall, 431 05-279 ambigua “Nuttall” Philippi, 448 05-310 ambiqua Weinkauff, 448 05-310 angulifera Lamarck, 455 05-329 antipodum Philippi, 471 05-355 arboricola Reeve, 459 05=333 ardouinianum Heude, 460 05-334 articulata “Menke” Philippi, 459 05-333 articulata Nevill, 464 05-338 articulata Philippi, 459 052333 australis Gray, 485 05-379 Austrolittorina Rosewater, 467 05-351 Bacalia H. & A. Adams, 431 05-279 Bacalia Gray, 431 05-279 balteata Reeve, 434 05-286 basterotii Payraudeau, 431 05-279 beccarii Tapparone-Canefri, 425 05-269 biangulata von Martens, 499 05-403 bicolor Lamarck, 519 05-455 blandfordi Dunker, 460 05-334 brevicula Philippi, 433 05-285 bullatus Martyn, 517 05-453 carinata Schrenck, 434 05-286 carinifera Menke, 464 05-338 carnifera Sherborn, 465 05-339 cecillei Philippi, 483 05-377 chaoi Yen, 493 05-397 Cidaris Réding, 513 05-449 cincta Quoy & Gaimard, 474 05-358 cinerea Pease, 505 05-409 cingulata Philippi, 459 05-333 coccinea Gmelin, 439 05-301 coccinea Martyn, 440 05-302 columna “Jonas” Philippi, 438 05-300 concolor Weinkauff, 438 460 conica Philippi, 465 contracta Nevill, 438 Corneolitorina Powell, 424 coronaria Lamarck, 517 coronatus Valenciennes, 515 crassior Philippi, 469 Cremnoconchus Blanford, 417 cumingi Philippi, 525 decollata Philippi, 477 delicatula Nevill, 460 diemensis Gray, 469 diemensis Quoy & Gaimard, 469 draparnauldii Audouin, 476 Echinella Swainson, 513 525 Echinellopsis Rovereto, 513 Echinininae Rosewater, 510 525 Echininus Clench & Abbott, 525 echinulatus Alth, 527 (Trochus) echinulatus Kiener, 527 elegans Philippi, 515 erronea Nevill, 460 493 eudeli Sowerby, 493 exigua Dunker, 500 Ezolittorina Habe, 431 Fectaria Philippi, 513 feejeensis Reeve, 500 fernandezensis Rosewater, 471 fijiensis “Reeve” Nevill, 500 flammea Philippi, 459 flammulata Philippi, 459 foliorum Rumphius, 459 fortunei Reeve, 459 fragilis Fenaux, 426 glabrata Philippi, 445 globulus Angas, 426 gracilior Philippi, 443 grandinatus Gmelin, 516 granicostata E.A. Smith, 493 granocostata Reeve, 492 granularis “Gray” of authors, 492 502 Granulilittorina Habe & Kosuge, 491 guttata Reeve, 476 [05-257] 05-300 05-334 05-339 05-300 05-268 05-453 05-451 05-353 05-261 05-467 05-361 05-334 05-353 05-353 05-360 05-449 05-467 05-449 05-444 05-467 05-467 05-469 05-469 05-451 05-334 05-397 05-397 05-404 05-279 05-449 05-404 05-355 05-404 05-333 05-333 05-333 05-333 05-270 05-307 05-270 05-305 05-452 05-397 05-396 05-396 05-406 05-395 05-360 532 Index Joseph Rosewater hagruma Tokioka & Habe, 493 Hamus “Klein” H. & A. Adams, 513 heterospiralis Grabau & King, 434 hisseyiana Tenison-Woods, 426 hybrids, 488 incisa Yokoyama, 466 529 indistincta Turton, 479 infans E.A. Smith, 452 insularis E.A. Smith, 493 intermedia Philippi, 459 Isonema “Hall” Provancher, 431 iwakiana Nomura & Hatai, 506 knysnaensis Philippi, 478 kowiensis Turton, 479 kozaiensis Nomura & Onisi, 453 kraussi Rosewater, 444 laevigata Schrenck, 434 Laevilacunaria Powell, 424 Laevilitorina Pfeffer, 424 laevior Nevill, 465 laevis Philippi, 443 laevis “Philippi” Reeve, 469 Lamellilitorina Tryon, 455 lamellosa Montrouzier, 426 leucosticta Philippi, 497 leucosticta “Philippi” Tryon, 499 limax Gray, 440 lineatum Gmelin, 459 Litonia Weinkauff, 431 Litorina Menke, 431 Litorinna Dall, 431 Litormia Morch, 431 Littoraria Griffith & Pidgeon, 435 littorea Linné, 431 Littorelaea Leach, 431 Littorina Férussac, 431 Littorina-capsula, 483 493 Littorinopsis Mérch, 455 Littorivaga Dall, 431 Littornia Jenkins & Grocock, 431 Littorrina Gabb, 431 luchuana Pilsbry, 528 lucida Yokoyama, 453 luctuosa Reeve, 474 lutea Philippi, 459 luteola Quoy & Gaimard, 459 05-397 05-449 05-286 05-270 05-382 05-340 05-317a 05-363 05-314 05-397 05-333 05-279 05-410 05-362 05-363 05-317 05-306 05-286 05-268 05-268 05-339 05-305 05-353 05-329 05-270 05-401 05-403 05-302 05-333 05-279 05-279 05-279 05-279 05-297 05-279 05-279 05-279 05-377 05-397 05-329 05-279 05-279 05-279 05-470 05-317 05-358 05-333 05-333 Littorinidae Macquariella Finlay, 424 major Swainson, 519 malaccana Philippi, 483 mandshurica Schrenck, 434 manschurica “Schrenck” Tryon, 434 marmorata Pfeiffer, 492 marmorata Philippi, 492 mauriciana “Lamarck” Potiez & Michaud, 443 mauritiana Lamarck, 442 melanacme E.A. Smith, 493 melanostoma Gray, 462 miliaris Quoy & Gaimard, 492 millegrana Philippi, 491 492, minor Weinkauff, 460 miodelicatula Oyama, 466 moerchi Adams & Angas, 426 monilifera Eydoux & Souleyet, 483 multistriata Tokioka, 483 natalensis Philippi, 489 Neritoides T. Brown, 431 Neritotrema “Recluz”” Wenz, 431 Neritrema Récluz, 431 newcombi Reeve, 459 newkombi “Reeve” in Weinkauff, 460 Nina Gray, 525 Nodilittorina von Martens, 481 nodosa Gray, 487 nodulosus Gmelin, 483 novaehiberniae Lesson, 459 novaezelandiae Reeve, 492 obesa Sowerby, 440 obtusata Linné, 431 oliveri Finlay, 471 opercula, 509 Pagodella Swainson, 513 Pagodus Gray, 513 pagodus Linné, 518 pallescens Philippi, 459 papillosa Lamarck, 521 papillosa of authors, 515 pascua Rosewater, 484 paytensis Philippi, 443 Pellilacunella Powell, 424 Pellilitorina Pfeffer, 425 penitaria Wood in G. Nevill, 472 perdix King & Broderip, 465 [05-258] 05-268 05-455 05-377 05-286 05-286 05-396 05-396 05-305 05-304 05-397 05-336 05-396 05-395 05-396 05-334 05-340 05-270 05-377 05-377 05-383 05-279 05-279 05-279 05-333 05-334 05-467 05-375 05-38 1 05-377 05-333 05-396 05-302 05-279 05-355 05-443 05-449 05-449 05-454 05-333 05-457 05-451 05-378 05-305 05-268 05-269 05-356 05-339 January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12 Index 533 perplexa Turton, 476 philippiana Habe & Kosuge, 493 philippiana Reeve, 459 philippina von Martens, 460 pica “Reeve” Turton, 479 picea Reeve, 479 picta Philippi, 503 pindata Philippi, 448 pintado Wood, 447 planaxis “Nuttall” Tryon, 504 praetermissa May, 445 Problitora Iredale, 426 pseudolaevis Nevill, 469 pulchella Dunker, 476 pultneyii Leach, 431 punctata Gmelin, 474 punctata Philippi, 459 pyramidalis Nevill, 465 pyramidalis Quoy & Gaimard, 481 483 quadriseriata Philippi, 515 radiata Eydoux & Souleyet, 492 rehderi Ladd, 524 reproduction of Tectarius, 510 reticulata Anton, 426 rhodea Biggs, 460 rietensis Turton, 479 Rissolittorina Ponder, 425 rubra Philippi, 459 rubropicta von Martens, 465 rugosa Menke, 487 rugosus Wood, 514 rusticus Philippi, 522 scabra Linné, 456 schmitti Bartsch & Rehder, 449 serialis Eydoux & Souleyet, 448 sieboldii Philippi, 459 sinensis Philippi, 459 sitchana Philippi, 431 songoense K. Martin, 523 souverbiana Crosse, 434 spinulosus Philippi, 527 05-360 05-397 05-333 05-334 05-363 05-363 05-407 05-310 05-309 05-408 05-307 05-270 05-353 05-360 05-279 05-358 05-333 05-339 05-375 05-377 05-451 05-396 05-460 05-444 05-270 05-334 05-363 05-269 05-333 05-339 05-381 05-450 05-458 05330 05-311 05-310 05-333 05-333 05-279 05-459 05-286 05-469 squalida Broderip & Sowerby, 431 strigata Lischke, 460 strigata Philippi, 459 strubelli von Martens, 493 subangulata Lamarck, 455 subcingulata Nevill, 460 Subditotectarius Ladd, 524 subgranosa Dunker, 499 subinermis Philippi, 523 subintermedia Nevill, 438 subnodosa Philippi, 495 suleatula Nevill, 438 sulcatus Nilsson, 431 sulculosa Philippi, 459 sundaica Altena, 450 suturalis Philippi, 459 syriaca Philippi, 476 Tectariinae Rosewater, 510 513 Tectarium, 523 Tectarius Valenciennes, 513 tectumpersicum Linné, 520 tenebrata “Nuttall” Jay, 448 tenuis Nevill, 460 tenuis Philippi, 438 trochiformis Dillwyn, 483 trochoides Gray, 483 tryphena Bartsch, 479 undulata Gray, 436 unifasciata Gray, 467 urieli Biggs, 493 ventricosa Philippi, 459 4992 vera Philippi, 521 verus J.E. Gray, 519 vidua Gould, 492 vilis “Menke” Philippi, 483 vitensis Dunker, 500 vitiensis “Reeve” von Martens, 500 vitrea Deshayes; 443 zebra Donovan, 435 zelandiae Finlay, 474 [05-259] 05-279 05-334 05-333 05-397 05-329 05-334 05-460 05-403 05-459 05-300 05-399 05-300 05-279 05-333 05-312 05-333 05-360 05-444 05-449 05-459 05-449 05-456 05-310 05-334 05-300 05-377 05-377 05-363 05-298 05-351 05-397 05-333 05-396 05-457 05-455 05-396 05-377 05-404 05-404 05-305 05-297 05-358 534 Index Joseph Rosewater Littorinidae Published by THE DEPARTMENT OF MOLLUSKS Delaware Museum of Natural History Box 3937, Greenville, Delaware 19807, U.S.A. [05-258] —— INDO-PACIFIC MOLLUSCA Monographs of the Marine Mollusks of the World with Emphasis on those of the Tropical Western Pacific and Indian Oceans EDITED BY R. TUCKER ABBOTT VOLUME 3 Published by DELAWARE MUSEUM OF NATURAL HISTORY Box 3937, Greenville, Delaware 19807, U.S.A. 74 Title Page Editors Volume 3 Notice Looseleaf subscribers should place this title page in the first binder immediately after the title page of volume 2. It will be your record of the number of volumes in your looseleaf set. [this sheet issued June 1, 1973, with vol. 3, no. 14] May 30, 1973 M\oll INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 1 WILLIAM H. DALL SECTIONAL LIBRARY DIVISION OF MOLLUSKS THE GENUS DRUPA IN THE INDO-PACIFIC by WituiaAM K. EMERSON Department of Living Invertebrates The American Museum of Natural History New York, N.Y. 10024 and WALTER O. CERNOHORSKY Auckland Institute and Museum Auckland, New Zealand Abstract A revised classification of the gastropod genus Drupa Roding (Muricidae: Thaidinae) is presented. The following taxa are recognized: Drupa (Drupa) morum morum Roding, 1798; D. (D.) morum iodostoma (Lesson, 1840); D. (D.) ricinus ricinus (Linnaeus, 1758); D. (D.) ricinus hadari Emerson and Cernohorsky, new subspecies; D. (D.) elegans (Broderip and Sowerby, 1829); D. (Ricinella) rubusidaeus Roding, 1798; D. (R.) speciosa (Dunker, 1867); D. (R.) clathrata clathrata| (Lamarck, 1816); D. (R.) clathrata miticula (Lamarck, 1822); D. (Drupina) grossularia Roding, 1798; and D. (Drupina) lobata (Blainville, 1832). Generic and specific synonymies are given for these taxa, together with distributional and ecolog- ical data for each species. Indo-Pacific Drupa Species of the genus Drupa Roding are con- fined in their distribution to the tropical Indo- Pacific region, where they are commonly en- countered on_ intertidal reef-flats. Drupa species are muricacean’ gastropods which show a close relationship with the larger, but otherwise similar and closely related species of Thais Roding, and are currently assigned to the subfamily Thaidinae within the family Muricidae. Because of the close similarity in shell morphology, species of Drupa Réding, and Morula Schumacher have frequently been considered to be congeneric or only subgener- ically separable. Wu (1965b), in his compara- tive study of the functional anatomy of the digestive systems of Drupa ricinus (Linnaeus) and Morula granulata (Duclos), found mor- phological differences in features of the rad- ula, gland-gut complex, stomach and _ rectal gland, and most notably in the structure of the salivary glands. The exterior of the animal of Drupa consists of a foot with an attached, chitinous oper- culum, a head, snout, proboscis, a pair of ten- tacles and eyes and the reproductive organ. Animals are dioecious, with the male’s penis situated behind the right tentacle below the thin mantle. The operculum is brown in color and corneous, stereotyped thaidine in appear- oe ae me Pe iS SOP . a OF Sad Plate 1. Camouflage in Drupa (Drupina) grossularia Réding. Cape Tuiolemu, Upolu, Samoa Ids. Top figure: Two specimens in situ on an exposed algae-covered reef. Bot- tom figure: two over-turned, living specimens on _ reef (photo courtesy A. Solem). [14-801] 2 Emerson and Cernohorsky Muricidae ance, with a series of plateaulike ridges on the occupying the mid-eulittoral zone. Demond side of attachment and concentric growth- rings on the exterior which converge basally into an ill-defined nucleus (see pl. 3, _ figs. 1-4). In the Fiji Islands and the New Hebrides, Drupa were encountered in the mid-eulittoral and upper eulittoral region of the intertidal zone, generally on windward and_ exposed reef-flats. Species were considerably less nu- merous on protected, coral-strewn or shingle- covered leeward reefs. Drupa morum Réding, D. rubusidaeus Réding, and D. ricinus (Lin- naeus) were most frequently collected on ex- posed algal ridges which were constantly kept moist through agitated waters and a_ breaking surf. The same species occurred on reef-edges covered with calcareous algae and_ detritus. Drupa ricinus, however, also occurred in the mid-eulittoral zone, while D. — grossularia Roéding, was usually confined to this part of the reef-zone. Kay (1971), in her study of the molluscan fauna of Fanning Island in the Line Islands, reported D. ricinus, D. morum and D. grossularia to be the most common macro- mollusks inhabiting reef-flats. D. ricinus and D. grossularia were among the dominant spe- cies of the beach-rock assemblages, with the former species restricted to exposed areas. D. ricinus also occurred on a_ subtidal, la- goonal patch reef, but was found to be far more abundant on seaward reefs than on the patch reefs in the lagoon. Salvat (1970), in his study of littoral molluscs of Fangataufa, Tua- motus, found a_ similar distributional pattern of Drupa as that observed by the junior author in Melanesia, with D. ricinus being more fre- quent on seaward algal ridges and D. morum Figs. 1-3. Drupa (Drupa) morum morum Réding, 1798. 1, Chisimaio, Somalia (ANSP 298192); 2, 3, adult and immature, both from Okinawa Id. (ANSP 302877). Figs. 4, 5. Drupa (Drupa) morum iodostoma (Lesson, 1840). Both from Ua Huka Id., Marquesas Ids. (ANSP 155617 and 156169). Figs. 6-8, 11. Drupa (Drupa) ricinus ricinus (Linnaeus, 1758). 6. Okinawa Id. (ANSP 302919); 7, 11, Mahé. Seychelles Ids. (ANSP 266229): 8, immature, Moorea, Society Ids. (ANSP 283222). Figs. 9, 10. Drupa (Drupa) ricinus new subspecies hadari Emerson and Cernohorsky. 9, paratype (AMNH 112617a); 10, holotype (AMNHII 166928). Eilat, Gulf of Aqaba, Israel. Fig. 12. Drupa (Drupa) elegans (Broderip and Sowerby, 1829). Society Islands (ANSP 199558). Figs. 13-15. Drupa (Ricinella) rubusidaeus Réding, 1798. 13, Gesira, Somalia (ANSP 299187); 14, Isles Radama, N.W. Madagascar (ANSP 257243); 15, immature; Malaita Id., British Solomon Ids. (ANSP 289624). (1957) reported D. grossularia as occurring in Micronesia most commonly on windward reef flats on rocks near the low tide line, less fre- quently on leeward reefs and rarely in la- goons. D. morum morum was recorded com- monly found living among rocks and coral of windward reef-flats and on windward _la- goonal reef-flats of the larger atolls, but rarely on leeward reefs. This species was encoun- tered most often near the low tide line, on or near the reef-edge, but also was taken in tide pools across the entire reef-flat. D. ricinus was reported to occupy a similar habitat to D. morum morum, while D. rubusidaeus was found to inhabit both windward and leeward reef-flats, living under rocks and coral rubble; it also occurred in tide-pools near, or just be- low the low tide line, and was found also liv- ing on coral heads off the seward reef-edge, in 10 to 15 feet of water. Heinicke (1970) en- countered D. lobata (Blainville), seemingly al- ways occurring in pairs, in the lagoonal chan- nel among coral heads at Diani Beach, Kenya. Drupa species are commonly encrusted with algae, coral growth, vermetids, Foraminifera, Hipponix and other’ extraneous’ organisms, making them blend in with the substrate upon which they rest (see pl. 1). Nothing is known about natural enemies of Drupa, except that Schoenberg (1971) records captive Conus tex- tile Linnaeus preying on Drupa morum, “D. speciosa” (= D. rubusidaeus), and both color forms of D. ricinus, among numerous other species of Hawaiian prosobranch gastropods. Conflicting reports may be found in litera- ture on the feeding habits of Drupa species. Salvat (1970) examined the microscopic con- Figs. 16-18. Drupa (Ricinella) clathrata clathrata (Lamarck, 1816). 16, Okinawa Id. (ANSP 289725); 17, Hivaoa Id., Marquesa Ids. (ANSP 155492); 18, Pacific Ocean (ANSP 36720). Figs. 19, 20. Drupa (Ricinella) clathrata miticula (Lamarck, 1822). 19, Arsenal Bay, Mauritius (ANSP 273087); 20, Mahébourg, Mauritius (AMNH 104995). Figs. 21, 22. Drupa (Ricinella) speciosa (Dunker, 1867). Both from “Rarotonga, Cook Ids.’—probably an error (ANSP 29873). Known from the Tuamotus and Pitcairn Islands. Figs. 23, 24. Drupa (Drupina) grossularia Roding, 1798. 23, Okinawa Id. (ANSP 225428); 24, immature, Sorsogon. Luzon Id., Philippines (ANSP 224148). Figs. 25, 26. Drupa (Drupina) lobata (Blainville, 1832). 25, Mogadiscio, Somalia (ANSP 295772); 26, Direction Id., Cocos-Keeling Ids., Indian Ocean (ANSP 288455). (all figures about natural size) [14-802] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 3 Plate 2. Genus Drupa Roding in the Indo-Pacific (all figures about natural size) Explanation on opposite page. [14-803] 4 Emerson and Cernohorsky tents of the digestive tracts of several reef- dwelling gastropods and decided that D. rici- nus and D. morum morum were herbivores, whereas other authors (Wu 1965b; Taylor 1968; Kay 1971; Cemohorsky, personal obser- vation) report these species to be carnivores. The feeding habits of the following species have been reported: Drupa morum morum worms and sipunculids (Kay, 1971) [Line Ids. ] barnacle _— Tetraclita 1968) [Seychelles Ids. ] herbivorous (Salvat, 1970) [Tuamotu Ids. } sipunculid worms (Cernohorsky, pers. ob- servation) [New Hebrides] Drupa ricinus live prey, i.e. sponges and_holothurians, or carrion (Wu, 1965b) [Hawaiian Ids. ] molluscs, barnacles and worms’ (Kay 1971) [Line Ids.] Drupa grossularia omnivorous (Salvat, 1970) [Tuamotu Ids. ] squamosa (Taylor, Plate 3. Opercula of Drupa. Figs. 1, 2. Drupa (Drupa) morum morum Réding, from Nananu-i-Ra Id., Fiji Ids. 1, inner surface; 2, outer surface. Fig. 3. Drupa (Ricinella) clathrata clathrata (Lamarck), from Pango Point, Efate Id., New Hebrides. Fig. 4. Drupa (Drupina) grossularia Réding, from Wadigi Id., Fiji Ids. Muricidae In comparison to Drupa_ species, Morula granulata (Duclos) was reported to feed on other mollusks and bamacles in the Seychelles Islands (Taylor, 1968), but was recorded prey- ing on holothurians, boring into bivalves of Isognomon and Ostrea, and consuming carrion in the Hawaiian Islands (Wu, 1965b). No information is recorded on the mode of reproduction of Drupa, although J. B. Taylor (in litt.) reports certain Hawaiian species of Drupa (sensu lato) to have planktonic veli- gers. Such a larval stage would account for the wide distribution of most species. Radulae The radula of Drupa is of the rachiglossate type, with 3 teeth per transverse row with a formula of 1-1-1. The radular ribbon is small and very narrow, and the lateral teeth are simple and sickle-shaped (see pl. 4, fig. B). The rachidian teeth are more or less subquad- rate or rectangular, the base is weakly con- cave, the central cusp is slender and longer than the flanking, bifid to quadrifid side- cusps. The lateral denticles are small, moder- ately deeply rooted and number from 2 to 5, and the end-cusps are usually slightly larger than the lateral denticles (see pl. 4, fig. A). Although the shells of Drupa show an af- finity with species of Morula Schumacher, the radulae of the moruline group of species dif- fer in the following particulars: the central cusp of the rachidian is more deeply rooted, the flanking side-cusps are not multifid as in Drupa, but are simple, and the central cusp and side-cusps are separated from each other by an interposing small, intermediate cusp. The radula of Morula is essentially muricine in appearance while that of Drupa is a weakly modified thaidine radula, which approaches that of Murex s. s. A classification based on radular morphol- ogy is complicated by the sporadic appear- ance of a drupine-type radula in other thai- dine genera, e.g. Agnewia tritoniformis (Blain- ville, 1832) [see Kesteven, 1902, pl. 29, fig. 5 and Cooke, 1919, text fig. 26], Semiricinula muricina (Blainville, 1832) [see Arakawa, 1965, pl. 14, figs. 19, 20], and Neothais smithi (Brazier, 1889). Wu (1965b) surmized that the distinctive drupine and moruline radula_pat- tern displayed by the species investigated by him may be directly associated with their re- spective feeding habits. [14-804] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 5 A Plate 4. Radula of Drupa (Drupa) ricinus ricinus (Linnaeus). One transverse row. Fig. A. rachidian or central tooth. Fig. B. lateral tooth. Abbreviations— ec, end cusps Id, lateral denticles sc, side cusps cc, central cusp Distribution and fossil record Species of Drupa are largely confined in distribution to the Indo-Pacific region, but some range from the northern regions of the Red Sea to Easter, Clipperton and the Gala~ pagos Islands in the eastern Pacific Ocean. They inhabit tropical waters and do not occur beyond latitudes of 35°N and 35°S. Although muricacean gastropods date from the _ late Mesozoic in the Cretaceous, typical thaid gas- tropods first appear in the Oligocene in mid- Tertiary (Keen, 1971). These Cenozoic forms apparently are the precursors of the drupine forms that are here referred to Drupa s.l. The drupine forms, with low-spired, — thaidlike shells having strong labial and columellar teeth, are poorly represented in the fossil record and are known only from the Pleisto- cene. Fossil moruline forms, with smaller, oval-biconical and higher-spired_ shells than those of Drupa s.l., are recorded as ranging from the Eocene to the Pleistocene (Eames, 1971, as “Drupa (Sistrum)”). The limited pale- ontological data suggest, therefore, that Drupa sl. and Morula_ s.l. evolved independently from a pre-thaid stock. The fossil evidence is not sufficient, however, to establish a_ well- documented evolutionary chronology of these generic groups. As pointed out by Hertlein (1960), Paleogene fossils from Europe and the East Indies have been placed in the genera “Ricinula” or “Sis- trum.” A survey of the literature indicates that some of these taxa are not closely allied to Drupa or Morula. Cossmann (1889, p. 132; 1903, p. 80, pl. 3, fig. 15) refers Purpura ringens Deshayes, 1865, from the Eocene of the Paris Basin to Ricinula, a spurious drupine assignment. Another example is Sistrum bay- lei Cossmann & Lambert, (1884, p. 175, pl. 5, fig. 19), from the Oligocene of France; both of these taxa appear to be buccinacean gastro- pods. Purpura styriaca Stur in Hilber, 1879, a Neogene fossil from the Miocene of Hungary, is a thaid, although this taxon was recently referred to Drupa by Strausz (1966, p. 284, fig. 130). Ricinula puruensis Martin (1914, p. 147, pl. 4, fig. 104) [not fig. 105], described from the late Eocene of Java, appears to be the earliest record for the genus Morula. Miocene species of Morula s.l. include: M. angsanana (Martin, 1921, p. 466, pl. 59, fig. 52), M. turrita (Martin, 1880, p. 41, pl. 8, fig. 3), both from Java, M. austriaca (Hoernes & Auinger, 1882, pl. 16, figs. 14-17) from the Vienna Basin, and M. in- constans (Michelotti, 1847, p. 217) from Italy. Neogene fossils that are purported to repre- sent living species of Morula s.l. are reported from Mio-Pliocene, Pliocene and_ Pleistocene deposits in the’ present Indo-Pacific faunal region and elsewhere. Drupa s.str. is recorded by Eames (1971) as ranging from the Pliocene to Recent and as occurring in the Red Sea, Indo-Pacific, East Africa) and questionably in North America. The New World record is based on the genus Condonia Hertlein, 1965, (type-species by orig- inal designation Sistrum hannai Howe, 1922), from the Pliocene of Oregon and California. This species, however, bears a resemblance to certain buccinacean shells, such as Can- tharus and Columbella, and the monotypic genus Condonia, therefore appears to be ref- erable to the superfamily Buccinacea instead of the Muricacea. Pentadactylus rhombiformis Martin (1899, p. 138, pl. 21, figs. 316a, b), described from the Pliocene of Java, was compared with sev- eral living muricacean species, including ref- erences to Reeve’s (1846, Purpura, pl. 3, fig. 13) illustration of Drupa rubusidaeus Roding. The species was considered closest in relation- ship to Purpura muricina Blainville, and the description and illustration of the Indonesian fossil suggest that it is a spinose thaid. Thus the available data indicate no valid records for Drupa s.l. prior to the Pleistocene. The following species are reported from Pleis- tocene deposits: Drupa (Drupa) morum Roding, 1798; D. (D.) ricinus ricinus (Linnaeus, [14-805] 1758), and Drupa (Ricinella) rubusidaeus Réd- ing, 1798 (see distributional records). Classification The Linnaean species of Drupa_ s.l. were originally described in the genus Murex Lin- naeus. Rodding (1798) proposed the genus Drupa; Montfort (1810) the genus Sistrum; and Lamarck (1816) the genus Ricinula, all for some of the drupine species previously assigned to Murex. The genus-group name Ricinula remained in use in malacological lit- erature until about 1913, when it was grad- ually replaced with Réding’s chronologically prior Drupa. Thiele (1929) accepted Drupa as a_ valid genus-group, but relegated several moruline genera, i.e. Cronia H. & A. Adams, Morulina Dall (=Azumamorula’ Emerson), — Phrygio- murex Dall, Maculotriton Dall and Drupella Thiele, as subgenera of Drupa. Wenz (1941) erected the new subfamily Drupinae, which besides the type-genus Drupa, contained the genus Thais R6ding and other thaid genera, together with the non-thaid genus Tritonalia Fleming (= Ocenebra Gray). Drupinae Wenz, 1938 and 1941 is presently considered a junior synonym of Thaidinae Suter, 1909 (as Thai- sidae Suter, 1909, Rec. Canterbury Mus., vol. 1, p. 11; 1909, Subantarctic Islands of New Zealand, art. 1, p. 27). The subfamilial name Thaidinae was conserved by action of the International Commission on Zoological No- menclature (Opinion 886, 1969) in preference to the chronologically older, but less fre- quently used Purpurinae. Species of Drupa are here assigned to the genus mainly on_ shell-morphology, although radular characters have also been considered. As pointed out in the section on “Radulae,” a classification on radular characters alone would require an inclusion of species of Ag- newia, Semiricinula, Neothais and a species of Morula. Such a classification was in fact pro- posed by Cooke (1919), who included 7 non- drupine species in Drupa on the basis of radular characters which he considered “dis- tinctly of the Drupa type.” It is obvious that on shell-morphology alone, the limits of Drupa are well-defined, but radular characters of Drupa also appear rarely in species refer- able to other thaidine genera. Due to the dispersal at auction of the Bolten collection, on whose specimens Réding’s new Emerson and Cernohorsky Muricidae descriptions were based, the whereabouts of the type-specimens are no longer known. F. C. Schmidt did purchase a small part of Bolten’s collection in 1819, but in a letter written by him, he observed that a great amount of the collection was purchased by Hamburg buyers. From those specimens procured by Schmidt from the Bolten collection, now in the Natur- kundemuseum, Staatliche Museen zu _ Gotha, Germany, only very few can be traced back to Bolten (Dr. Motschmann, in litt.). In the ab- sence of Rd6ding’s type-specimens we have designated appropriate cited illustrations of specimens figured by other authors as lecto- types of Roding’s species. List of Recognized Taxa Below are listed the recognized generic and specific taxa for the genus Drupa. The eleven species and subspecies are referred to three genus-groups. All are living, and three are also recorded as Pleistocene fossils. Family Muricidae Rafinesque, 1815 Subfamily Thaidinae Suter, 1909 Genus Drupa Réding, 1798 Subgenus Drupa Réding, 1798 morum morum Roding, 1798. Type species. Recent. Indo-Pacific. and Eastern Pacific, except the Marquesas Islands. Pleistocene. morum iodostoma (Lesson, 1840). Recent, Marquesas Islands. ricinus ricinus (Linnaeus, 1758). Recent, Indo-Pacific and Eastern Pacific. Pleisto- cene. ricinus hadari Emerson and _ Cernohorsky, new subspecies. Recent, Red Sea. elegans (Broderip and Sowerby, 1829). Re- cent, Wake Island to the Tuamotu Islands. Subgenus Ricinella Schumacher, 1817 rubusidaeus R6ding, 1798. Type species. Re- cent, Indo-Pacific. Pleistocene. speciosa (Dunker, 1867). Recent, Tuamotu and Pitcairn Islands. clathrata clathrata (Lamarck, 1816). Recent, tropical west Pacific Ocean. clathrata miticula (Lamarck, 1822). Recent, Indian Ocean. Subgenus Drupina Dall, 1923 grossularia Roding, 1798. Type species. Re- cent, East Indian Ocean and Pacific. lobata_ (Blainville, 1832). Recent, Ocean. Indian [14-806] May 30, 1973 Abbreviations The following institutional abbreviations are used in this paper: AIM—Auckland Institute and Museum, Auck- land AMNH—American Museum of Natural History, New York AMS— Australian Museum, Sydney ANSP—Academy of Natural Sciences of Phil- adelphia BM (NH)—British Museum (Natural History), London BPBM— Bernice P. Bishop Museum, Honolulu DM—Dominion Museum, Wellington DMNH—Delaware Museum of Natural History, Greenville FMNH—Field Museum of Natural History, Chi- cago MCZ—Museum of Comparative Zoology, Cam- bridge, Massachusetts LACMNH—Los Angeles County Museum of Natural History MHNG—Museum d Histoire Naturelle, Geneva SDMNH—San Diego Museum of Natural His- tory USNM—National Museum of Natural History, Washington, D.C. WAM—Western Australian Museum, Perth Acknowledgments We gratefully acknowledge the help ex- tended to us in providing access to collections, technical assistance, loan of specimens, field data and information on types. We would like to thank the following persons: R. T. Abbott—DMNH; E. Binder—MHNG; W. J. Clench and R. D. Turner—MCZ; S. P. Dance, National Museum of Wales, Cardiff; R. K. Dell—DM; the late A. Hadar, Tel Aviv, Israel; E. A. Kay, University of Hawaii, Hono- lulu; J. Knudsen, University Zoological Mu- seum, Copenhagen; Y. Kondo—BPBM,; Fei-Jann Lin, Academia Sinica, Taipei, Taiwan; J. H. McLean—LACMNH,; D. F. McMichael—formerly AMS; D. Motschmann, Naturkundemuseum, Gotha; W. E. Old, Jr.—AMNH; V. Orr Maes— ANSP; A. W. B. Powell—AIM; H. A. Rehder, J. Rosewater and J. P. E. Morrison—USNM; G. E. Radwin—SDMNH; Mme. P. Revercé, Nou- mea, New Caledonia; M. G. Richards—formerly AMNH:; B. Salvat, Muséum National d’Histoire Naturelle, Paris; V. Siewersten, Koloa, Hawaii; A. Solem—FMNH; Mr. & Mrs. G. D. Stout, INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 7 New York; J. B. Taylor, Prescott College, Ari- zona; J. Taylor and K. Way—BM (NH); N. Teb- ble, Oxford University Museum; J. J. Wage- man, Koloa, Hawaii; C. S. Weaver, Kailua, Hawaii; B. R. Wilson and S. M. Slack-Smith— WAM; Shi-Kuei Wu, University of Michigan, Ann Arbor. Species excluded from Drupa Included under this heading are species of Thaidinae which were originally described in Drupa, or have been referred to this genus by subsequent authors. In the latter category, only those species requiring further explana- tion have been listed. Neothais bollonsi (Suter, 1906) (Pls. 5, 6) Remarks—Suter (1909) assigned this mor- uline species to the genus Drupa on the basis of the typically drupine radula. Iredale (1915) correctly synonymized D. bollonsi with the southeast Australian species Purpura smithi Brazier, 1889, and assigned it to the genus Neothais Iredale, 1912. Despite its drupine radular characters, the species should be placed near Morula Schumacher. amas aan / Md hat i Olam Plate 5. Radula of Neothais smithi (Brazier). Half a trans- verse row; Sunday Id., Kermadec Ids. [synonym is Drupa bollonsi Suter]. Plate 6. Neothais smithi (Brazier). [synonym is Drupa_ bol- lonsi Suter]. Fig. 1. Sunday Id., Kermadec Ids. (AIM; 24.5 x 17.4 mm). Fig. 2. Norfolk Id. (AIM; 18.0 x 12.0 mm). [14-807] 8 Emerson and Cernohorsky Synonymy— 1889 Purpura smithi Brazier, Australian Museum Memoir, no. 2, p. 28, pl. 4, figs. 1-4, 7-12, 21-22 (Lord Howe Id.) [as Purpura (Polytropa) smithi on plate ex- planation]. 1902 Purpura tritoniformis var. smithi Brazier, Kesteven, Proceedings of the Linnaean Society of New South Wales, pt. 4, p. 534. 1906 Purpura striata Martyn subsp. bollonsi Suter, Trans- actions and Proceedings New Zealand Institute, vol. 38, p. 331 (Kermadec Ids.). 1909 Drupa bollonsi Suter, Proceedings of the Malacolog- ical Society of London, vol. 8, p. 254, pl. 11, figs. 5-7 (shell, operculum and_ radula); 1913 Suter, Manual of the New Zealand Mollusca, p. 428, pl. 19, fig. 11 (New Zealand). 1915 Neothais smithi (Brazier), Iredale, Transactions and Proceedings New Zealand Institute, vol. 47, pp. 474, 475; 1915 Oliver, Transactions and Proceed- ings New Zealand Institute, vol. 47, p. 536; 1950 Dell, Dominion Museum Records, Zoology, vol. 1, no. 3, p. 26. Condonia hannai (Howe, 1922) Remarks—This moderately large, (66.8 mm long) species from the Pliocene of Oregon and California’ was originally described in the genus Sistrum Montfort. Hertlein (1965) pro- posed for this species the new genus Con- donia, and placed the genus in the muricid subfamily “Drupinae” with apparent reluc- tance. He concluded that: “Condonia hannai bears a general resemblance to the Recent Drupa iodostoma Lesson but the spire of that species is low and the columella [=error for inner margin of outer lip] bears denticles typical of Drupa.” He also noted that the genus was not known as a fossil in the Eastern Pacific region, but that living representatives occur in the Galapagos Islands and at Clipper- ton Island. Although the labial dentition does superficially resemble that of Drupa, the type- species of Condonia lacks columellar den- ticles. This extinct species appears to be a buc- cinacean gastropod, and Dr. G. E. Radwin (in litt.) considers Condonia hannai to be an extralimital representative of the genus Col- umbella s.str. which, perhaps due to its un- usual northern, cooler-water habitat, attained a giant size. The monotypic genus Condonia, therefore, appears to be referable to the Buc- cinacea rather than the Muricacea. Synonymy— 1922 Sistrum hannai Howe, Univ. California Publ., Bull. Dept. Geol. Sci., vol. 14, no. 3, p. 102, pl. 8, figs. 1, 5 (Fossil Point, S. W. Empire City, Coos Bay, Plio- cene of Oregon); 1943 Weaver, Univ. Washington Publ. Geology, vol. 5, pt. 2, p. 450, pl. 87, figs. 14, Muricidae 16 (figured holotype); 1960 Hertlein, Veliger, vol. 3, no. 1, p. 8 (San Benito County, Pliocene of Cali- fornia). 1965 Condonia hannai (Howe), Hertlein, Occas. Papers California Acad. Sciences no. 49, p. 4, figs. 3, 4 (figured holotype). Azumamorula mutica (Lamarck, 1816) (Pls. 7, 8) Remarks—For the western Indian Ocean spe- cies Ricinula mutica Lamarck, which is inter- mediate in shell-characters between Drupa Roding, and Morula Schumacher, Dall (1923) proposed the genus-group Morulina. Thiele (1929) and Wenz (1941) assigned Morulina as a subgenus to Drupa. Emerson (1968) pro- posed the substitute name Azumamorula for the preoccupied Morulina Dall (non Morulina Borner, 1906, in Insecta), and figured the rad- ula of the type-species, which is typically moru- line. Plate 7. Radula of Azumamorula mutica (Lamarck). Half a transverse row; Black River Bay, Mauritius (after Azuma and d’Attilio in Emerson, 1968). Plate 8. Azumamorula mutica (Lamarck). Fig. 1. Holotype from unknown locality (“Mozambique” on label) [MHNG no. 1101/19; 20.2 x 16.5 mm]. Synonymy— 1816 Ricinula mutica Lamarck, Tabl. Encycl. Methodique, p. 1, pl. 395, figs. 2a, b (no locality given); 1846 Reeve, Conchologia Iconica, vol. 3, pl. 2, fig. 1. 1822 Ricinula pisolina Lamarck, Hist. nat. anim. s. ver- tebres, vol. 7, p. 233 (Ile de France - Mauritius); 1835 Kiener, Spec. gen. icon. coquilles vivantes, vol. 8, p. 20, pl. 4, fig. 8a (juvenile specimen); 1844. Deshayes and Milne-Edwards, Hist. Nat. anim. s. vertebres, ed. 2, vol. 10, p. 52. 1919 Morula mutica (Lamarck), Cooke, Proceedings of the Malacological Society of London, vol. 13, p. 106 (description of radula). [14-808] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 9 1923 Morulina mutica Lamarck, Dall, Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 75, p. 303. 1929 Drupa (Morulina) mutica (Lamarck), Thiele, Handb. syst. Weichtierkunde, vol. 1, p. 294; 1941 Wenz, Handb. Palaozoologie, vol. 6, pt. 5, p. 1112, text fig. 3159. 1968 Azumamorula mutica ‘Lamarck), Emerson, Nautilus, vol. 81, no. 4, p. 125, text fig. (radula). Drupa vitiensis Pilsbry in Pilsbry and Bryan, 1918 Remarks—The species illustrated by Pilsbry and Bryan is the male form of Drupella cornus (Roding, 1798). Generally credited to Pilsbry, 1921, the specific name was validated by Pils- bry and Bryan through a published illustration in combination with a specific name. Synonymy— 1918 Drupa vitiensis Pilsbry in Pilsbry & Bryan, Nautilus, vol. 31, no. 3, pl. 9, fig. 5. 1921 Sistrum’ vitiense Pilsbry, Proceedings of the Acad- emy of Natural Sciences of Philadelphia, vol. 72, p. 319 (Fiji Ids.). Drupa walkerae Pilsbry and Bryan, 1918 Remarks—The species described by the au- thors belongs to Drupella Thiele, 1925, and is similar to, if not conspecific with D. rugosa (Born, 1778). Synonymy— 1918 Drupa walkerae Pilsbry & Bryan, Nautilus, vol. 31, no. 3, p. 99, pl. 9, fig. 4 (Honolulu Harbor, Hawai- ian Ids.). 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B., vol. 254, pp. 129-206, pls. 13-17, text figs. Tomlin, J. R. le B. 1934. The marine Mollusca of Christ- mas Island, Indian Ocean. Bulletin Raffles Museum, Singapore, vol. 9, pp. 74-84. Wilson, B. R. and K. Gillett. 1971. Australian Shells. A. H. & A. W. Reed, Sydney, 168 pp., 106 pls., text figs. Wu, S-K. 1965a. Studies of the radulae of Taiwan muricid Gastropods. Bulletin Institute Zoology, Acad- emia Sinica, vol. 4, pp. 95-106, 35 text figs. Wu, S.-K. 1965b. Comparative functional studies of the digestive system of the muricid gastropods Drupa ricina and Morula granulata. Malacologia, vol. 3, no. 2, pp. 211-233, pls. 1-5. Gastro- [14-811] Shout tie ail on Walter O. Cernohorsky has been a malacolo- gist and the Curator of Molluscs at the Auckland Institute and Museum, Auckland, New Zealand, since 1969. Born in Czecholovakia on June 30, 1927, he emigrated to Fiji in the 1950’s where he was Chief Surveyor for the Emperor Gold Mining Company and served as Honorary Conchologist at the Fiji Museum. In 1968, he received a senior post-doctoral Research Associateship with the Smithsonian Institution, Washington, D. C. He is author of numerous scientific papers on the marine mollusks of the Indo-Pacific, especially those of the families Mitridae, Muricidae and Strombidae. His contributions have appeared in The Nautilus, The Veliger, the Bulletin of the Auckland Institute and Museum, and the Revue Suisse de Zoologie. In 1967 and 1972 he authored two volumes for amateurs entitled, Marine Shells of the Pacific. Mr. Cernohorsky is an accom- plished photographer, an active field collector, is married, and has two children. 2 Emerson and Cernohorsky Muricidae buh Dr. William K. Emerson is Curator of Mollusks and Chairman of the Department of Living In- vertebrates at the American Museum of Natural History, New York City. His research interests concern taxonomy and zoogeography of late Cen- ozoic marine mollusks, especially gastropods and scaphopods. He has participated in several ex- peditions to west Mexico and the Caribbean region. A native of California, Dr. Emerson was born in San Diego on May 1, 1925, and earned degrees in zoology from the California State University at San Diego (A.B. 1948) and the University of Southern California, Los Angeles (M.S. 1950). From 1951 to 1955, he served as Museum Paleontologist at the University of Cali- fornia, Berkeley, where he received a Ph.D. in invertebrate paleontology in 1956. He joined the staff of the American Museum in 1955. Dr. Emerson is the author of numerous scientific articles and is co-author of three books designed to assist amateurs. He is a past president of the American Malacological Union and of the Wes- tern Society of Malacologists. [14-812] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 13 Family Muricidae Rafinesque, 1815 Subfamily Thaidinae Suter, 1909 Key To Subgeneric Groups of Indo-Pacific Drupa A. Columella singularly folded axially B. Labial teeth compound, constricting the aperture Gash duike vats ankle ena tee Drupa s.str. p. 14-819 BB. Labial teeth singularly arranged or occasionally in close association, not compound, aperture not constricted Prt arenes rete eee ae ar Ricinella p. 14-841 AA. Columella doubly folded axially; margin of outer lip with 2 digitate processes, aperture narrow, NOt CoONStricted ........ cee eeeceeeeeeceeeee ees Subfamily Thaidinae Suter, 1909 The subfamily name Thaidinae has been granted precedence over the long established family-group name Purpuridae Menke, 1828, in Opinion 886 of the International Commis- sion on Zoological Nomenclature. Although the Commission has dated Thaidinae Suter from 1913, Suter erected the family-group name in 2 prior publications in 1909 (see syn- onymy). Authors generally divide the family Muri- cidae into 7. subfamilies: the Muricinae, Ocenebrinae, Aspellinae, Thaidinae, Trophon- inae, Typhinae and Rapaninae. Some authors, however, consider the Thaidinae worthy of family rank, but since no scientific evidence as to important anatomical, morphological or ecological differences between the Thaidinae and other closely related muricid groups has been presented, an elevation of Thaidinae to family rank appears unwarranted. In such a closely related species-group as the Muricidae, with numerous existing species of ‘interme- diate generic characters, a taxonomic consoli- dation rather than further subdivision, is indi- cated. err Srey oer Te Drupina p. 14-859 Apart from the nominate type-genus Thais Réding, 1798, the subfamily Thaidinae_ con- tains such genera as Mancinella Link, 1807, Acanthina Fischer von Waldheim, 1807, Cymia Morch, 1860, Purpura Bruguiére, 1789, Nucella Réding, 1798, Nassa Réding, 1798, Vexilla Swainson, 1840, Pinaxia H. & A. Adams, 1853, Xanthochorus Fischer, 1888, Morula Schu- macher, 1817, Azumamorula Emerson, 1968, Drupella Thiele, 1925, Neothais Iredale, 1912, Lepsiella Iredale, 1912, Agnewia Tenison- Woods, 1878, Cronia H. & A. Adams, 1853, and other subgeneric groups and fossil genera. Synonymy— 1828 Purpuracea Menke, Synopsis methodica Mollus- corum, p. 34 (suppressed by the ICZN in Opinion 886, 1969, Bull. zool. Nomenclature, vol.’ 26, pp. 128-132). 1839 Purpuridae Broderip, Penny Cyclop., vol. 14, p. 321 (suppressed in Opinion 886 of ICZN). 1840 Purpurinae Swainson, Treatise on Malacology, p. 71 (suppressed in Opinion 886 of ICZN). 1909 Thaisidae Suter, Records Canterbury Museum, vol. 1, p. 11; 1909 Suter, Subantarctic Islands of New Zealand, art. 1, p. 27. 1938 Drupinae Wenz, Handbuch der Palaozoologie, vol 6, pt. 1, pp. 42, 47; 1941 Wenz, ibid., pt. 9, p. 1112. [14-817] 14 Emerson and Cernohorsky Subfamily Thaidinae Suter, 1909 Genus Drupa Roding, 1798 Type: Drupa morum Réding, 1798 On shell characters, the group generally considered as belonging to Drupa s.str. can be divided into two distinct groups. In the group of the type-species Drupa morum_ Réding, which also includes the subspecies D. morum iodostoma (Lesson), D. ricinus (Linnaeus) and D. elegans (Broderip & Sowerby), the majority of the denticles on the outer lip are arranged as compound, i.e. bifid or trifid teeth which constrict the aperture posteriorly. In the group comprising D. rubusidaeus Réding, D. speci- osa (Dunker) and D. clathrata (Lamarck), the denticles of the outer lip are singularly situ- ated as non-compound teeth, which results in an appreciably wider aperture. For this group of species the subgeneric name Ricinella Schumacher, 1817, is available. Although the teeth on the outer lip of Drupa (Drupina) grossularia Réding, the type-species of the subgenus Drupina Dall, are also singularly arranged, the shell of the species develops marginal lobate processes and the radula_ has a greatly modified rachidian radular tooth and very slender and small lateral teeth. Drupa differs from Morula in having a more sub-ovate form, lower spire, longer aperture and a_ structurally different central radular tooth. Members of Drupa show a close radular relationship with most of the thaidine groups. Muricidae Synonymy— 1798 Drupa Réding, Museum Boltenianum, p. 55. Type- species by subsequent designation, Rovereto, 1899: Drupa morum Réding, 1798. 1807 Canrena Link, Beschr. Nat.-Samml. Univ. Rostock, p. 126. Type-species by monotypy: Canrena_neri- toidea Link, 1807 [= Drupa morum Roding, 1798] (as restricted by Iredale, 1937, and lectotype. des- ignation by Cernohorsky, 1969). 1810 Sistrum Montfort, Conchyliologie systématique, vol. 2, p. 595. Type-species by original designation: Sistrum album Montfort, 1810 [= Drupa_ ricinus (Linnaeus, 1758) ]. 1816 Ricinula’ Lamarck, Tableau Encyclopédique — et Methodique, p. 1. Type-species by subsequent designation, Children, 1823: Ricinula horida (= R.horrida Lamarck) 1798]. 1822 “Ricinella Lam.”, Bowdich, Elements of Conchology, vol. 1, p. 40. [? error for Ricinula Lamarck, 1816] (sole species listed and figured Ricinula horrida Lamarck, 1816) [non Ricinella | Schumacher, 1816]. 1852 Pentadactylus Mérch, Catal. Conchyl. Yoldi, vol. 1, p. 87. Type-species by subsequent designation, Baker, 1895: Pentadactylus ricinus Lamarck = Drupa ricinus (Linnaeus, 1758). [non — Pentadactylus Schultze, 1760, in Echinodermata; nec Gray, 1845, in Reptilia]. 1855 Ricimula Gould, U.S. Astronom. Exp. South. Hemi- sphere, vol. 2, p. 263 (error for Ricinula Lamarck, 1516). 1859 Ricinulus Demarest in Chenu, Encycl. Hist. Nat. Crust. Moll. Zooph., p. 174 (invalid emendation for Ricinula Lamarck, 1816). As pointed out by the junior author (Cerno- horsky, 1969), Suter (1913) is credited by Dodge (1957) and other authors with the type designation of Drupa, but Rovereto’s (1899) designation is earlier. Although members of _ this [= Drupa morum Rodding, genus were placed until the turn of the present century in Ricinula, Lamarck’s there were available Plate 9. Type-species of subgenera of Drupa Réding. _ Fig. 1. Drupa (Drupa) morum Réding. Namoui reef, Niue Id. (WOC coll.; 29.8 x 27.4 mm). Fig. 2. Drupa (Ricinella) rubusidaeus Riding. Suva reef, Fiji Ids. (WOC coll.; 40.5 x 36.5 mm). Fig. 3. Drupa (Drupina) grossularia Réding. Pango Point, Efate Id., New Hebrides (WOC coll.; 28.3 x 28.6 mm). [14-818] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 15 three valid prior generic names, of which Drupa Roding, 1798, is the oldest. Link, 1807, established Canrena as a monotypic genus with Canrena neritoidea Link, the sole species. His citation to Martini’s figures include the species Drupa ricinus (Linnaeus), D. morum Réding, and D. grossularia Réding. Canrena neritoidea was restricted by Iredale (1937) to Drupa morum Roding, as depicted by Mar- tinis figures 972, 973, on plate 101. The same figures were designated as the lectotype of the species Canrena neritoidea Link, by Cern- ohorsky (1969). The monotypic genus Sistrum Montfort, 1810, has been erroneously used by many authors for the species rightly contained in Morula Schumacher, 1817, which has Drupa uva Réding, as the type-species. Sistrum al- bum Montfort, the type-species of Sistrum by original designation, is a synonym of Drupa ricinus (Linnaeus). In his discussion of Sistrum album, Montfort considered the species to have a white, violet or yellowish aperture, a misconception common with writers of the day. Although the violet aperture would indi- cate the species Drupa morum, Montfort’s fig- ure of Sistrum album, the specific name itself and its French, Dutch and Flemish equivalents of “Le Sistre blanc,” “La mure blanche,” “wite moerbesie” and “witte moerbeyer,” all refer to the white-mouthed form of Drupa ricinus (Linnaeus). Subgenus Drupa sensu stricto Shell small to medium, strong, heavy, sub- ovate; whorls low or conical, often with si- phonous tubercles, 2 tubercles rarely extended from the margin as lobate processes; surface generally sculptured with imbricated — scales; aperture noticeably constricted posteriorly by labial teeth; teeth arranged in compound groups; columella typically with 1 prominent axial fold, rarely with 2 folds, and with 3-5 prominent oblique plications; radular formula 1-1-1, lateral teeth fang-like, similar in shape, central tooth typically composed of a large medial cusp, flanked by a pair of slightly smaller, bifid to quadrifid cusps, and small, slender lateral denticles; operculum sub-lunar to sub-linear, nucleus marginal, marginal cal- lus well developed, muscle scars few in num- ber, arranged as gyratory lines. Members of this genus are typical coral and reef forms, being confined to the tropical waters of the Indo-Pacific region, although some species occur at Easter, Clipperton and the Galapagos Islands, in the tropical eastern Pacific. Drupa morum morum Roding, 1798 (PI. 2, figs. 1-3; Pls. 10, 11) Range—Red Sea and East Africa to the East- ern Pacific including Easter and Clipperton Is- lands (except the Marquesas Islands). Remarks—The_ purple-colored aperture and the shorter tubercles distinguish this species from the white- or yellow-mouthed form of D. ricinus. The possession of tubercles, the deeper purple color of the aperture and the lack of dorsal bands offset this from the Mar- quesan subspecies D. morum iodostoma (Les- son, 1840). Habitat—Intertidal on reef-flats, among rocks and in crevices; frequently found near the reef-edge on reefs exposed to strong surf. Description—Shell 18 to 49 mm (4% to 2 inches) in length, ovately globose, spire acu- minate, generally inconspicuous’ in adult stage. Whorls nodose with four rows of short, strong tubercles. Interstices between tubercles striated with scalelike striae. Aperture — nar- row, linear, extending nearly three-quarter the length of the shell. Outer lip crenulated be- tween tubercles, dentate on the interior mar- gin with plaitlike teeth; teeth arranged in an upper group of four denticles and in a lower group of three denticles. Two conspicuous plaits situated immediately above the deep canal. Inner lip enameled with a callus con- taining 3 or 4 plaitlike ridges projecting into the aperture above the canal. Posterior si- phonal canal elongate, obliquely recurved to- ward the apex. Color white or grayish, tuber- cles dark-brown; interior of aperture deep purple in adults, light purple in immature specimens. Operculum typical of the group. Naa O..mm Plate 10. Radula of Drupa (Drupa) morum morum Roding. Half a transverse row; Fiji Ids. [14-819] 16 Drupa The central cusp of the rachidian of the radula is slightly longer than the flanking and bifid side-cusps; the side-cusps are followed by 3 to 5 small lateral denticles, exclusive of the slightly stronger end-cusps. In some speci- mens, the lateral denticles descend onto the side-cusps. Measurements (mm)—(including specimens with a mature lip). spines; all length width 49.1 45.9 S. E. Zanzibar 40.0 38.2 Lectotype of horrida Lamarck 32.4 29.6 Tosa, Japan 27.0 23.7 Viti Levu, Fiji 18.1 17.1 Bikini Atoll, Marshalls Synonymy— 1684 Buonanni, Rec. ment. oculi anim. test., (3), fig. 173 only (very poor). 1685 Lister, Hist. Syn. Meth. Conchyliorum, pl. 804, fig. 13. 1753 Klein, Tent. meth. ostr. nat. cochl., pl. 1, fig. 30. 1758 Nerita nodosa Linnaeus (pars), Systema Naturae, ed. 10, p. 777 (refers to Lister, Klein and Buon- anni, (fig. 173) only). 1767 Murex neritoideus Linnaeus (pars), Systema Na- turae, ed. 12, p. 1219 (refers to Lister, Klein and Buonanni (fig. 173) only; 1791 Gmelin (pars), Sys- tema Naturae, ed. 13, p. 3537 (refers to Lister (fig. 13 only), Buonanni, Klein, Knorr and Martini (figs. 972, 973 only) and Seba (fig. 41 only); 1825 Wood, Index Testaceologicus, p. 123, pl. 26, fig. 47a. Knorr, Verg. Augen u.Gemiuths, pt. 1, pl. 25, figs. 5, 6 (good). 1777 “Murex Morum_ globosum” Martini, Syst. Conch- ylien-Cabinet, vol. 3, p. 280, pl. 101, figs. 972, 973 (East Indies and coast of Coromandel) [non_ bi- nomial]. 1798 Drupa morum Roding, Museum Boltenianum, p. 55 refers to Martini and Knorr) [no locality given]; 1913 Hedley, Nautilus, vol. 27, no. 7, p. 80; 1936 Hirase, Coll. Jap. shells, p. 79, pl. 110, fig. 8; 1957 Kaicher, Indo-Pacific Sea Shells, pl. 4, fig. 3; 1960 Hertlein, Veliger, vol. 3, no. 1, p. 8 (Gala- pagos and Clipperton Ids.); 1961 Rippingale & Mc- Michael, Queensland and Gt. Barrier reef shells, pl. 13, fig. 5; 1965 Arakawa, Venus: Jap. Journ. Malacology, vol. 24, no. 2, p. 114, pl. 13, figs. 3, 4 (radula); 1965 Wu, Bull. Inst. Zool. Acad. Sinica, vol. 4, p. 98, text fig. 19 (radula); 1967 Orr Maes, Proc. Acad. Nat. Sci. Philadelphia, vol. 119, no. 4, p. 129; 1969 Cernohorsky, Veliger, vol. 11, no. 4, p. 298, pl. 47, fig. 7 (shell), text fig. 4 (radula); 1970 Salvat, Cahiers du Pacific, no. 14, p. 46; 1971 Wilson & Gillett, Australian Shells, p. 92, pl. 61, fig. 2; 1971 Kay, Pacific Science, vol. 25, pp. 266, 275. 1807 Canrena_ neritoidea Link, Beschr. Nat.-Samml. Univ. Rostock, 3 Abth., p. 126 (refers to Martini, pl. 101, figs. 972, 973—designated as lectotype figures by Cernohorsky, 1969) [no locality given]. 1816 Ricinula horrida Lamarck, Tabl. Encycl. Methodique, p. 1, pl. 395, figs. la, b (no locality given); 1822 Lamarck, Hist. nat. anim. s. vertebres, vol. 7, p. 231 (Indian Ocean); 1823 Sowerby, Genera Rec. foss. shells, pt. 18, pl. 235, fig. 1; 1842 Reeve, Conchologia Systematica, vol. 2, p. 215, pl. 156, fig. 1; 1846 Reeve, Conchologia Iconica, vol. 3, pl. 1, fig. 3; 1859 Chenu, Manuel Conchyliologie, 1768 Emerson and Cernohorsky Muricidae vol. 1, p. 168, fig. 814; 1880 Tryon, Manual Con- chology, vol. 2, p. 184, pl. 56, figs. 201, 202; 1933 Dautzenberg & Bouge, Journal de Conchyliologie, vol. 77, p. 238. 1817 Ricinella violacea Schumacher, Essai Nouv. Sys- téme, p. 240 (refers to Martini, op. cit.) [no local- ity given]. 1823 Ricinula horida (sic) Children, Quart. Journ. Sci. Lit. & Arts, vol. 16, p. 56, pl. 5, fig. 189. 1832 Purpura horrida Lamarck, Blainville, Nouv. Ann. Mus. d’Hist. Nat. Paris, ser. 3, vol. 1, p. 208; 1833 Quoy & Gaimard, Voyage L’Astrolabe, vol. 2, p. 576, pl. 39, figs. 1-3 (animal and operculum); 1835 Kiener, Spéc. gen. icon. coq. viv., vol. 8, p. 8, pl. 1, fig. 1 (animal). 1850 Sistrum horridum M. Gray, Figs. Moll. Animals, vol. 4, p. 70, pl. 96, fig. 11; 1911 Schepman, Si- boga-Expeditie, vol. 49d, p. 355; 1952 Morris, Field Guide shells Pacific coast and Hawaii, pl. 39, fig. 5. 1852 Ricinula (Pentadactylus) globosa Mart., Morch, _ Cat. Conchyl. Yoldi, vol. 1, p. 88 (synonymized with R. horrida Lamarck and Drupa morum Roding). 1853 Pentadactylus (Pentadactylus) globosus H. & A. Adams, Genera Rec. Mollusca, vol. 1, pp. 129, 130, pl. 13, fig. 6 (animal). 1908 Pentadactylus (Pentadactylus) horridus | Lamarck, Horst & Schepman, Cat. Syst. Moll. Mus. Hist. Nat. Pays-Bas, vol. 13, p. 157. 1938 Drupa (Drupa) morum Roding, Adam & Leloup, Mém. Mus. Roy. d’Hist. Nat. Belg., vol. 2, fasc. 19, p. 164. Types—Since Roding’s types are probably no longer extant, we designate the specimen de- picted on plate 101, figs. 972, 973 in Martini, as the lectotype of Drupa morum Roding (pl. 11 fig. 1). Two syntypes of Ricinula horrida Lamarck, are in the Muséum d Histoire Natur- elle, Geneva; the specimen measuring 40.0 mm in length, no. 1101/12/1, is here selected as the lectotype of R. horrida. The type-speci- men of Ricinella violacea Schumacher, could not be located in the University Zoological Museum, Copenhagen. From the two localities mentioned by Martini for D. morum s. s. we select the first-mentioned “East Indies” as the Plate 11. Drupa (Drupa) morum morum Roding. Fig. 1. Lectotype figure from Martini, 1777, Syst. Con- chylien-Cabinet, vol. 3, pl. 101, fig. 972. Fig. 2. Lectotype of Ricinula horrida Lamarck (MHNG no. 1101/12/1; 40.0 x 38.2 mm). [14-820] May 30, 1973 type locality, and restrict it further to Java, Indonesia (specimens in AMNH and ANSP). Nomenclature—This species was well known to early naturalists but was confused by Lin- naeus with Thais nodosa and T. nodosa ascen- sionis Quoy & Gaimard (= meretricula Roding). In his 12th edition of the “Systema Naturae,” Linnaeus re-named the species Murex neri- toideus, and cited the same erroneous figures which depict Drupa morum. However, in Lin- naeus’ personal copy of the 12th edition of the “Systema Naturae,” the complete synonymy of the 10th edition description of Nerita nodosa has been cancelled and the following words were added: “Labium interius punitis 2 maculatum.” Records—RED SEA: Lahonel Beach, Gulf of Aqaba, Israel (Lamy, 1938, p. 56); Berenice, Egypt (ANSP); Mah- mud reef, Jiddah, Saudi Arabia (AMNH). EAST AFRICA: SE Point, Isla di Serpenti, Chisimaio, Somalia (AMNH); Pangavini Id., 10 mi. NNE_ Dar-es-Salaam, Tanzania (MCZ); Diani Beach, Kenya (Heinicke, 1970, p. 7); Moz- ambique (USNM). ZANZIBAR: Chumbe Id.; Kiwengwa; Ras Nungwe (all ANSP). SEYCHELLES: (AMNH); Cous- in Id. (ANSP). MADAGASCAR: S. side of Nossi Iranja, 32 mi. SW of Nossi Bé; Flacourt, Fort Dauphin; 2 mi. NE Point Ansiraikiraiky (all MCZ). MAURITIUS: 1 mi. ESE of Souillac; Pointe Fayette (both ANSP); near Port Louis (MCZ); Point Pimente, N. side Arsenal Bay (Powell coll.). REUNION ISLAND: (ANSP). CHAGOS ARCHIPELAGO: (Melvill, 1909, p. 104). MALDIVE ISLANDS: Imma Id., SE North Male Atoll; Fadiffolu Atoll; Tiladummati Atoll (all ANSP), LACCADIVE ISLANDS: (Hornell, 1922, p. 217). INDIA: Pamban and Shingle Id., Gulf of Manaar (Satya- murti, 1952, p. 160). CEYLON: 12 mi. N. of Trincomalee (AMNH); Hikkaduwa (ANSP). THAILAND: Loam Seng, S. of Laam Son, Phuket Id. (ANSP). VIETNAM: Cam Ranh Bay (ANSP). COCOS-KEELING ISLANDS: N. end of Pulo Siput; West Id., Cocos Id. (both ANSP). CHRISTMAS ISLAND: (Tomlin, 1935, p. 79). INDONESIA: Jesselton district, N. Borneo; Pulan Boenta, off Acheh Head, N. W. Sumatra; Sulau Bay, Batu group, off Sumatra; Mega, Mentawai Ids., S. : W.Sumatra (all USNM); Queen’s Bay, INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 17 off Sukabumi, Java (AMNH); Keledjitan, Bantam, Java (ANSP); Batjan Id.; Manipa Id., W. of Halmahera Id._; Mandidi Id.; Ambayana Id., all Moluccas (all MCZ). PHILIPPINES: Corregidor Id.; Manila Bay, Luzon Id.; Calapan, Mindoro Id.; Gigmoto, Catanduanes Id.; Cuyo Id., Palawan group (all ANSP); Panay; Silino Id., Minda- nao Id. (both USNM); Sanga Sanga Id., Sulu Archipelago (ANSP). FORMOSA (Taiwan): (AMNH); Suo; Karenko; Botan-wan; Ryukyu-Syo; Hoko (all Kuroda, 1941, p. 111). RYUKYU ISLANDS: Hyakuna reef (AMNH); Yomitan reef, Okinawa: Bolo Point; Kuzu-Saki, all Okinawa (all ANSP); NE coast of Iheya Shima (ANSP); Miyako (FMNH). JA- PAN: Kikaiga Shima, Osumi; Oshima Osumi (MCZ; USNM); Tosa (ANSP); Kii, Honshu (FMNH). MARIANAS: Maung: Agrihan (both USNM); Saipan; Tinian; Guam (all AMNH). PALAU ISLANDS: Babelthuap (ANSP); Koror; Eil Malk (both USNM). CAROLINE ISLANDS: Kayangel; Angulpelu; Gorokoru; Ngargersiul (all ANSP); Yap (USNM); UWlithi; Ifalik; Elato; Satawal; Ponape; Kaping- amarangi (all ANSP). MARSHALL ISLANDS; Eniwetok Atoll; Rongelap Atoll; Majuro Atoll; Ujae Atoll; Uterik Atoll (all USNM):; Bikini Atoll; Arno Atoll (both AMNEFHI). WAKE ISLAND: (AMNH; ANSP; MCZ). NEW GUINEA: Schouten Ids.; Wewak:; Makiri reef, Unea Id. (all ANSP); 2 mi N. of Gusika, 13 mi. N. of Finschhafen (MCZ). AD- MIRALTY ISLANDS: Koruniat Id. (ANSP). AUSTRALIA: Queensland: Herald Cay, Coral Sea; Pipon reef near Cape Melville (both AMS): Wilson Id., Capricorn group (MCZ); Lady Elliot Id. (ANSP; AIM); Heron Id. (AIM); Tryon Id., Capricorn group (Powell coll.); | Northern Territory: Dar- win (FMNII); New South Wales: Middleton reef, off coast of New South Wales (Iredale, 1937, p. 256); Lord Howe Id. (AMS): West Australia: Barrow Id. (Wilson & Gillett, 1971, p. 92). NEW BRITAIN: Rabaul Harbour (ANSP). NEW IRELAND: Kavieng (AMNH). SOLOMON ISLANDS: NW. Bougainville Id. (ANSP); Lutee, Choiseul Id.; Ataa, N. Malaita) Id.; Lunga, Guadalcanal; Bellona Id. (all AMNIIL); Vikopia Id. (AIM); Bougainville; Bumana (both Powell coll). NEW) HEBRIDES: Tongoa, Espiritu) Santo Id. (MCZ); Bushmen’s Bay, E. Malekula Id. (AIM); Tanna Id. (DM). NEW CALEDONIA: Touho (AMNII); Isle of Pines (MCZ). LOYALTY ISLANDS: Lifu) (USNM). FIJI ISLANDS: Savusavu, Vanua Levu (AMNH); Korolevu, S. Viti Levu (ANSP); Nananu-i-Ra Id., N. Viti Levu (WOC coll.); Ongea Levu, Lau Ids. (USNM). GILBERT IS- LANDS: Onotoa Atoll (MCZ); Kingsmill Id. (USNM); Nauru Id. (Iedley, 1903, p. 4). ELLICE ISLANDS: Funa- futi (AMS: AIM). WALLIS ISLANDS: Nukuhifala (USNM). TONGA | Islands: Hlufagalupe, Tongatapu (ANSP); Ni- uafo’ou Id. (USNM). SAMOA ISLANDS: Asau_ Harbour, 160° 180° 160° T 140° \20° 100° 90° T r T Plate 12. Geographical distribution of Drupa (Drupa) morum morum Réding, and its Marquesan subspecies D. (D.) (Lesson). Open circles are literature morum iodostoma records. (14-821] 18 Drupa Savaii; Pago Pago, Tutuila (both USNM); Apia, Upolu (AMNH; AIM). NIUE ISLAND: Alofi (USNM); Oneone reef; Namoui reef (both AMNH). PHOENIX ISLANDS: Canton Id. (ANSP). COOK ISLANDS: Bird’s Id.; North Id., both Palmerston Atoll (both USNM); Aitutaki; Mauke; Rarotonga; Mangaia (all USNM); Kopuano Passage, Aitu- taki (AIM). PENRHYN ISLAND: (AMNH; AIM). AUSTRAL ISLANDS: Tubuai; Raivavae; Rurutu (all USNM). SO- CIETY ISLANDS: many localities on Bora Bora; Huahine; Moorea; Tahiti (all USNM); Venus Point, Tahiti (AMNH). TUAMOTU ISLANDS: Tikahau Atoll; Vahitahi; Fakarava; Nengonengo; Makatea (all USNM). GAMBIER ISLANDS: Mangareva (USNM). LINE ISLANDS: Palmyra _ Id. (USNM):; Fanning Id. (AMS); Caroline Id.; Flint Id. (both ANSP); Jarvis Id. (Powell coll.); Kingman reef; Washing- ton Id.; Christmas Id. (all Kay, 1971, p. 275). HAWAIIAN ISLANDS: Kamilo, Kauai; Hanauma Bay, Oahu; Pukoo, Molokai; Honokowai reef, Maui; Hilo, Hawaii (all USNM); Kona, Hawaii (AMNH); Koko Head, Oahu; Waikiki, Hon- olulu (both Powell coll.). PITCAIRN ISLAND: (AMNH). EASTER ISLAND: (Steele, 1957, p. 112). CLIPPERTON ISLAND: (Hertlein & Allison, 1960, p. 15). Fossil records—HAWAIAN ISLANDS: Pleistocene: Oahu (Ostergaard, 1928, p. 6; Kosuge, 1969, p. 786, pl. 5, fig. 93). Drupa morum iodostoma (Lesson, 1840) (PI. 2, figs. 4, 5; Pl. 13) Range— Marquesas Islands. Remarks—The lack of tubercles, the more quadrate shape, the dark dorsal bands and the pinkish violet aperture distinguish this uncom- mon subspecies from the nominate species Drupa morum Roding. Early records from both literature and spec- imens are from numerous localities in the western Indo-Pacific, but reliable data indi- cate this taxon to be restricted to the Mar- quesa Islands. Records based on old _ collec- tions in the National Museum of Natural His- tory, Washington, and the Museum of Com- parative Zoology, Harvard, can be confidently dismissed as being based on erroneous local- ity data, probably copies from locality indica- tions given in old literature; these localities are “Malacca,” “New Zealand,” “Madagascar” and “Fiji Islands.” However, there are several records of “D. iodostoma” from neighbouring areas to the Marquesas, such as Palmyra Island in the Line Islands (SDNHM), Venus Point, Tahiti (AMNH) and Fakarawa Atoll, Tuamo- tus (ANSP); in addition, the subspecies has also been reported in literature from Marutea Island (Couturier, 1907) and Makatea Island (Boettger, 1918), both in the Tuamotu Archi- pelago. The literature records are suspected Emerson and Cernohorsky Muricidae misidentifications, while the actual specimens in the mentioned institutions have either been obtained through secondhand or have been documented by persons who also collected in the Marquesas Islands apart from the Tuamo- tus. Confirmed, recent collections suggest, therefore, that D. iodostoma is an allopatric subspecies of D. morum, and is endemic to the Marquesa Islands where D. morum is not known to occur. Habitat—In surge channels cut into exposed volcanic shores (G. D. Stout, personal com- munication), and in rocky crevices at a depth of 10 feet. Description—Shell 32 to 47 mm (about 1% to 2 inches) in length, quadrately globose, spire very short and acuminate. Whorls faintly ax- ially plicate, spirally ribbed by five wide, slightly raised ribs which are more __pro- nounced toward the margin. Interstices be- tween ribs striated with 3 lirations. Aperture narrow, linear, extending nearly three-quarter the length of the shell. Outer lip crenulated between ribs, inner margin dentate; teeth compound, 3 to 4 denticles in the upper tooth, 2 to 3 denticles in the lower tooth. Above the anterior canal are two conspicuous plaits. Col- umella with three to four heavy plications pro- jecting into the aperture. Posterior siphonal canal elongate, obliquely curved toward the apex. Color cream, spiral ribs brownish black, intersticial spiral threads reddish brown; aper- ture pinky-violet. The radula is similar to Drupa morum; the rachidian has bifid side-cusps and 4 or 5 lat- eral denticles. The radula of “Drupa_ iodos- toma” as figured by Cooke (1919) from the “Sandwich Islands” [= Hawaiian Ids.], is probably based on Hawaiian specimens of D. morum. Measurements (mm)—(all specimens with a mature lip). length width 47.0 43.0 Nukuhiva, Marquesas Ids. 38.0 36.5 Nukuhiva, Marquesas Ids. 36.0 32.7 Tahuata, Marquesas Ids. 30.3 27.0 Tahuata, Marquesas Ids. Synonymy— 1840 Purpura (Ricinula) iodostoma Lesson, Rev. Zool. Soc. Cuvierienne, vol. 3, p. 355 (“New Zealand” = error!). 1842 Purpura iodostoma Lesson, Guerin’s Magasin de Zoologie, vol. 4, p. 58, pl. 58. [14-822] May 30, 1973 LO Plate 13. Radula of Drupa (Drupa) morum iodostoma (Les- son). Half a transverse row; Tahuata, Marquesa Ids. 1846 Ricinula iodostoma Lesson, Reeve, Conchologia Iconica, vol. 3, pl. 1, figs. 4a, b; 1860 Reeve, Ele- ments of Conchology, vol. 1, p. 82, pl. 7, fig. 32; 1880 Tryon, Manual of Conchology, vol. 2, p. 184, pl. 56, fig. 199. 1888 Ricinula jodostoma (sic) Lesson, Paetel, Catalog der Conchyl.-Sammlung, vol. 1, p. 143 (New Zealand [erroneous] ). 1908 Pentadactylus (Pentadactylus) iodostomus Lesson, Horst & Schepman, Cat. Syst. Moll. Mus. Hist. Nat. Pays-Bas, vol. 13, p. 154. 1913 Drupa iodostoma Lesson, Hedley, The Nautilus, vol. 27, no. 7, p. 80; 1957 Kaicher, Indo-Pacific Sea Shells, pl. 4, fig. 5 (Malaysia and Melanesia [= error!]); 1965 Hertlein, Occ. Papers California Academy of Sciences, no. 49, p. 2, figs. 1, 2 (Mar- quesas Islands). 1918 Ricinula jodostoma_ (sic) Lesson, Boettger, Abh. Senckenb. Naturfr. Gesellschaft, vol. 36, (3), p. 298 [Probably misidentified Drupa morum Rod- ing]. Types—The type-specimen of Drupa_ iodos- toma (Lesson) is probably in the Muséum Na- tional d’Histoire Naturelle, Paris. The given locality “New Zealand” is erroneous, as_ the species does not live there. Since the type- specimen was collected during the voyage of the “Vénus,” which visited the Marquesas Phi 0.1mm LSA 5.0 mm Plate 14. Drupa (Drupa) ricinus ricinus (Linnaeus)—white mouthed form; Fiji Islands. Fig. A. Radula; half a transverse row. Fib. B. Part of rachidian of radula showing variation in the number of accessory denticles on the side-cusps. Fig. C. Penis. INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 19 after the 14th of August, 1838, and also called at the Bay of Islands, New Zealand, during October 1838 (Chamberlin, 1960, p. 67), the erroneous locality is obviously due to a mix-up of specimens during the voyage. Since reliable records are known only from the Marquesas, we designate Taiohae, Nukuhiva Island, Mar- quesa Islands, as the type locality of D. morum iodostoma. Records—MARQUESAS ISLANDS: Hana Nui and Haavie Bays, Ua Huka Id.; Eiao Id. (all ANSP); Taiohae, Nuku- hiva Id. (USNM; AMNH); N. side Hana Moe Noe Bay, Tahuata (USNM). Drupa ricinus ricinus (Linnaeus*, 1758) (Pl. 2, figs. 6-5, 11; Pls. 14, 15, 16) Range—From East and South Africa to the Eastern Pacific, at Easter, Clipperton and Gal- apagos Islands. Remarks—Differs from D. morum, which has a proportionally larger shell, in having longer spines, a smaller size and in lacking the pur- ple or mauve apertural coloration. D. elegans (Broderip & Sowerby) differs mainly by _ pos- sessing a continuous, red-brown line — encir- cling the aperture; it is restricted to Polynesia where it is found with the present species. °It is the editorial policy to allow authors to use their own preference in spelling this name. Linnaeus or Linne. 0.1mm B Plate 15. Drupa (Drupa) ricinus ricinus (Linnaeus)—yellow mouthed form; Fiji Islands. Fig. A. Radula; half a transverse row. Fig. B. Part of rachidian of radula showing variation in the side-cusps and lateral denticles. [14-823] 20 Drupa Commonly, D. ricinus may have a faint orange or yellow, diffused line encircling the aper- ture. Populations with large shells, attaining nearly 40 mm in length, are present in the Red Sea. This form appears to be geographically isolated from the smaller-shelled, nominate subspecies, and it is recognized herein as a new subspecies Drupa ricinus hadari. Habitat—Lives exposed, or in water up to approximately 10 feet deep, on or under rocks, sand, weed or coral, on exposed wave benches or fringing reefs. At Clipperton Island the species has been collected to a depth of 22 fathoms. Description—Shell 17 to 35 mm (about % to 1% inches) in length, solid, obovoid, spire short, reduced, obscure in adult specimens, body whorl three-quarter the length of the shell. Whorls ribbed with five rows of sharp spines; spines variable in development, but higher near the margin of the aperture. Inter- stices between spines striated with scale-like striae. Aperture linear, constricted by the teeth and callus. Apertural dentition as in D. morum with features less prominent. Color white or grayish; spines dark-brown to dark- gray, especially toward the tips; mouth white or with a wide, broken, diffused, orange to yellow line extending around the outer lip and neighbouring canal area. Operculum typical for the group. In the radula, the side-cusps of the rachid- ian are short or moderately long, bifid to quadrifid, and are followed by 2-3. side- denticles, exclusive of the end-cusps. Measurements (mm)—(including spines; all) specimens With a mature lip). length width 32.3 27.8 Kagoshima, Japan 29.5 25.5 Lectotype of arachnoides Lamarck 25.6 27.0 S. Viti Levu, Fiji Ids. 23.0 25.0 Probable holotype of ricinus Linnaeus 22.8 21.0 Penrhyn Id. 17.2 16.3 Samarai, Papua Synonymy— 1705 Rumphius, Amboinsche Rariteitkammer, pl. 24, fig. E. 1742 d’Argenville, L’Histoire Naturelle ...., pl. 17, fig. A. 1742 Gualtieri, Ind. test. Conchyliorum, pl. 28, fig. N (poor). Emerson and Cernohorsky Muricidae 1758 Murex ricinus Linnaeus, Systema Naturae, ed. 10, p. 750 (refers to Rumphius and Gualtieri, op. cit.) [Asiatic Ocean] (yellow and white forms). 1758 Murex hystrix Linnaeus, Systema Naturae, ed. 10, p. 750 (refers to d’Argenville, op. cit.) [no locality given] (juvenile specimen); 1855 Hanley, Ipsa Linn. Conch., p. 294. 777 “Murex Morum globosum” Martini, Syst. Conchyl- ien-Cabinet, vol. 3, p. 280, pl. 102, figs. 976, 977 (non binomial) [yellow form]. 1798 Drupa tribulus Roding, Museum Boltenianum, p. 55 (refers to Rumphius and Gualtieri, op. cit.) [no locality given]. 1798 Drupa_ rubuscaesius Roding, ibid., p. 55 (refers to Martini, op. cit.) [no locality given] (yellow form). 1810 Sistrum album Montfort, Conchyliologie Systemat- ique, vol. 2, p. 595, fig. on p. 594 (white form) [no locality given]. 1816 Ricinula arachnoides Lamarck, Tableau Encyclo- pedique Méthodique, p. 1, pl. 395, figs. 3a, b (no locality given) [yellow form]; 1822 Lamarck, Hist. nat. anim. s. vertebres, vol. 7, p. 232 (Indian Ocean); 1823 Sowerby, Gen. Rec. fossil shells, pt. 15, pl. 235, fig. 5 (juvenile specimen); 1842 Reeve, Conchologia Systematica, vol. 2, p. 215, pl. 256, fig. 5 (juvenile specimen); 1846 Reeve, Con- chologia Iconica, vol. 3, pl. 1, fig. 5; 1859 Chenu, Manuel de Conchyliologie, vol. 1, p. 168, fig. 812. 1831 Murex neritoideus Mawe, Wodarch’s Intr. Conchol- ogy, pl. 3, fig. 43 (yellow form) [non Linnaeus, 1767]. 1832 Purpura albo-labris Blainville, Nouv. Ann. d Hist. Nat. Paris, ser. 3, vol. 1, p. 208, pl. 9, fig. 5 (Trin- comalee, Ceylon) [white form]; 1835 Kiener, Spéc. gen. icon. coq. viv., vol. 8, p. 12, pl. 1, fig. 2. 1832 Purpura arachnoides Blainville, ibid., p. 209 (yel- low form); 1833 Quoy & Gaimard, Voyage L’Astro- labe, vol. 2, p. 579, pl. 39, figs. 17-19 (animal and operculum); 1835 Kiener, Spéc. gén. icon. coq. viv., vol. 8, p. 10, pl. 1, figs. 3, 3a; 1848 Krauss, Stida- frik. Mollusken, p. 115 (Natal). 1850 Sistrum arachnoides Lamarck, M. Gray, Figs. Moll. Animals, vol. 4, p. 70, pl. 96, fig. 2. 1853 Pentadactylus ricinus Linnaeus, H. & A. Adams, Gen. Rec. Mollusca, vol. 1, p. 130; 1875 Troschel, Gebiss der Schnecken, vol. 2, p. 134, pl. 13, fig. 5 (radula). 1859 Ricinula albolabris Blainville, Chenu, Manuel de Conchyliologie, vol. 1, p. 168, fig. 812. 1880 Ricinula ricinus Linnaeus, Tryon, Manual Conchol- ogy, vol. 2, p. 184, pl. 56, fig. 200 and pl. 57, figs. 204, 206, 212; 1933 Dautzenberg & Bouge, Journal de Conchyliologie, vol. 77, p. 240. 1884 Pentadactylus arachnoides Lamarck, Fischer, Man- uel de Conchyliologie, fasc. 7, p. 646, pl. 6, fig. 9. 1911 Sistrum ricinus Linné, Schepman, Siboga-Expeditie, vol. 49d, p. 354. 1913 Drupa ricinus Linne, Hedley, Nautilus, vol. 27, no. 7, p. 80; 1937 Hertlein, Proc. Americ. Phil. Society, vol. 78, no. 2, p. 308, pl. 1, figs. 5, 6 (Clipperton and Galapagos Ids.); 1967 Orr Maes, Proc. Acad. Nat. Sci. Philadelphia, vol. 119, no. 4, p. 129; 1969 Cernohorsky, Veliger, vol. 11, no. 4, p. 299, pl. 47, figs. 8, 8a (shell), text figs. 5, 6 (radula); 1970 Salvat, Cahiers du Pacifique, no. 14, p. 46; 1971 Kay, Pacific Science, vol. 25, pp. 266, 275; 1971 Wilson & Gillett, Australian Shells, p. 92, pl. 61, figs. 3, 3a. 1915 Drupa rubus-cestus Dall, Smithsonian Inst. Publ. no. 2360, p. 29 (refers to Réding, 1798, p. 55, species 695 = D. tribulus Roding). 1929 Drupa (Drupa) ricinus Linnaeus, Thiele, Handb. syst. Weichtierkunde, vol. 1, p. 295; 1938 Adam & [14-824] May 30, 1972 Leloup, Mém. Mus. Roy. d’Hist. Nat. Belg., vol. 2, fasc. 19, p. 164. 1933 Ricinula ricinus var. arachnoides Lamarck, Dautzen- berg & Bouge, Journal de Conchyliologie, vol. 77, p. 240. 1960 Drupa ricina Linnaeus, Hertlein, Veliger, vol. 3, no. 1, p. 8; 1965 Arakawa, Venus: Jap. Journ. Malacol- ogy, vol. 24, no. 2, p. 115, pl. 13, fig. 1 (radula); 1965 Wu, Bull. Inst. Zool. Acad. Sinica, vol. 4, p. 98, text fig. 18 (radula); 1965 Wu, Malacologia, vol. 3, no. 2, p. 211, text figs. (anatomy). 1960 Drupa_ ricina forma albolabris Blainville, Hertlein, Veliger, vol. 3, no. 1, p. 8. 1965 Drupa albolabris (Blainville), Arakawa, Venus: Jap. Journ. Malacology, vol. 24, no. 2, p. 114, pl. 13, fig. 2 (radula). 1965 Drupa arachnoides Lamarck), Wu, Bull. Inst. Zool. Acad. Sinica, vol. 4, p. 98, text fig. 18 (radula). Types—The probable — type-specimen _ of Drupa ricinus (Linnaeus) [Pl. 16, fig. 2], is in the Linnean collection of the Linnean Society, London. This particular specimen is the yel- low-spotted form, which has a no. 540 written on the columella (the number of Murex ricinus in the 12th edition of the “Systema Naturae’”). Two additional, undocumented specimens are also in the collection, an adult of the white form and a juvenile specimen. Five syntypes of Ricinula arachnoides Lamarck, are in the Muséum d'Histoire Naturelle, Geneva, and the 29.5mm long syntype, no. 1101/15/4, is here selected as the lectotype (PI. 16 fig. 4). The type-specimen of Purpura albolabris Blain- ville, is probably in the Muséum _ National @Histoire Naturelle, Paris. The type locality of D. ricinus is “Asiatic Ocean,” which is here restricted to Ceylon (specimens in MCZ, ANSP and AMNH). Nomenclature—Much_ confusion has existed regarding the identity of this taxon. Lamarck, 1822, considered Murex hystrix Linnaeus, to be the pink-apertured species which is correctly known as Drupa_ rubusidaeus Roding. This misconception was continued by many early authors. Actually, as is pointed out by Hanley (1855), Linnaeus referred Regenfuss’ (1758) excellent colored figure of Drupa rubusidaeus Réding to Murex hippocastanum in his 12th edition of the “Systema Naturae,” and not to Murex hystrix. In the 10th edition, the short diagnosis of Murex hystrix was accompanied by only one figure reference, pl. 17, fig. A of d’Argenville, 1742. This engraving (see Pl. 16 fig. 1) unmistakeably represents the dorsal as- pect of Drupa ricinus as indicated by the long, obliquely slanted spines which characterize some forms of this species. In the description Linnaeus states: “apertura edentula_ repanda,” INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa All Plate 16. Drupa (Drupa) ricinus ricinus (Linnaeus). Fig. 1. Lectotype figure of Murex hystrix Linnaeus, from d’Argenville, 1742, L’Histoire Naturelle . Conchyli- ologie, pl. 17, fig. A. Fig. 2. Probable type-specimen of Murex ricinus Linnaeus, from the “Asiatic Ocean” (Linnean Society of London coll.; 23.0 x 25.0 mm). Fig. 3. Lectotype figure of Drupa rubuscaesius Réding, from Martini, 1777, Syst. Conchylien-Cabinet, vol. 3, pl. 102, tig. 976. Fig. 4. Lectotype of Ricinula arachnoides Lamarck (MHNG no. 1101/15/4; 25.5 x 29.5 mm). an indication that this species was based on specimens in which the dentate lip has not yet developed. Thus it can be safely assumed that Murex hystrix Linnaeus, is nothing more than a juvenile of Drupa ricinus Linnaeus. At the present time, some students recognize two species, the form with a white aperture as Drupa albolabris (Blainville) and the yellow- orange spotted form as D. arachnoides (Lam- arck). The latter should not be confused with the yellow-mouthed D. grossularia Réding. In a de- tailed sampling of D. ricinus in the Fiji Islands by the junior author (Cernohorsky, 1969), both [14-825] 22 Drupa forms were found not only to be sympatric in var- ious localities, but were sharing the same rock in many instances. No sexual dimorphism was ob- served in either color form, and no differences in either living animal, radula, color of the stomach pouch or the penis were apparent in either form. The number of accessory side-denticles in the rachidian of the radula varied from 3 to 5 in both color forms. Both color forms of D. ricinus have been collected together at various other Indo- Pacific localities. Hertlein and Allison (1960) re- corded the presence of both color forms of D. ricinus at Clipperton Island, an isolated coral atoll in the eastern Pacific, located about 670 miles southwest of Acapulco, Mexico. The white- apertured form was reported to be living among boulders and coral debris on the outer parts of the reef flats, and off the edge of the reef flats, in coral and coral rubble, to a depth of at least 130 feet, the lower limit of collecting by their SCUBA divers. Although the white-apertured form was found in abundance, only abraded specimens of the yellow-orange form were on the beach and off shore at a depth of 70 feet. In the New Hebrides the white-mouthed form only was found by the junior author. Records—(inserted in brackets: y = yellow-spotted form, w = white form). GULF OF ADEN: Aden (Shopland, 1896, p. 220). EAST AFRICA: Isla di Serpenti, Chisimaio, Somalia (AMNH; yw); Port Amelia, Mozambique (AMNH; y); Kend- wa Id., 4 mi. ESE Dar-es-Salaam, Tanzania; Pangavini Id., 10 mi. NNE Dar-es-Salaam, Tanzania; Ras Kankadya, 6 mi. N. Dar-es-Salaam, Tanzania (all MCZ; y); Diani Beach, Kenya (AMNH; USNM; y); Mombasa, Kenya (AMNH; y). ZANZIBAR: (AMNH; w); Ras Nyngwe; Chumbe Id.; Jembi- ani, 5 mi. S. Paje; Kiwenga (all ANSP; y). SOUTH AFRICA: Umtwalnmi, 22 mi. N. Port Shepstone, Natal; East London (both ANSP; y). SEYCHELLES: Loraie Bay, Curiense Id.; Anse aux Pins (both ANSP; y); Coetivy Id.; Praslin Id. (Mel- zo” 40° 60° 80° to0° zoe t40e Emerson and Cernohorsky Muricidae vill, 1909, p. 103). MADAGASCAR: Ambodifototra, Isle St. Marie (MCZ; w); N. E. of Pointe Antsiraikiraiky, NW Isle St. Marie; Ambariobe, S.E. Nossi Bé; Grande Recife, Tulear; Pointe Ibanona, Fort Dauphin (all MCZ; y). REUNION: (ANSP; w). MAURITIUS: Pointe Fayette; S.W. Port Louis (both ANSP; w); 1 mi. NW Black River (ANSP; y). MAL- DIVE ISLANDS: Ongu Id., N. Malosmadulu Atoll; Wala Id., Nilandu Atoll; Fadifolu Atoll (all ANSP; y); Imma Id., N. Male Atoll (ANSP; w). CEYLON: (MCZ; w); Pointe de Gal- lett (ANSP; y); 12 mi. N. of Trincomalee (AMNH,; y). ANDA- MAN ISLANDS: (Melvill & Sykes, 1899, p. 222) THAI- LAND: Laam Seng, 1 mi. S. Laam Son, Phuket Id. (ANSP; y); Goh Huyong, Similan Ids. (USNM; y). VIETNAM: Con Son Ids. (Fischer, 1891, p._ 149). COCOS-KEELING IS- LANDS: Direction Id. (ANSF; y); E. side of Horsburgh Id.; SW side West Id. (both ANSP; y). CHRISTMAS ISLAND: (Tomlin, 1934, p. 79). INDONESIA: Pulau Bai, Batu group, off Sumatra; Pulau Stupai, Mentawai Ids., S.W. Sumatra; Pelaboean Ratoe, Preanger, Java (all USNM; y); Morotai Id., Moluccas (MCZ; y); Mantanani Id. and Mandi Darrah Id., N. Borneo (both ANSP; y); Batu Dua and Palau Pombo, Wasi, Ambon I. (both FMNH; y); Malawali, N. Borneo (AMNH; y). PHILIPPINES: Many localities throughout the Archi- pelago; Luzon Id.; Mindoro Id.; Cebu Id.; Sulu Archipelago (AMNH; ANSP; y); Samar Id.; Catanduanes Id.; Palawan Id. (AMNH; ANSP; yw); Calamianes group; Mindanao Id. AMNH: ANSP; MCZ; w); Borongan village, Samar Id.; Gig- moto, Catanduanes Id. (both AIM; yw). FORMOSA (Taiwan): Tainan beach (AMNH; w); Suo; Kasyo-to; Botanwan; Ryukyu-syo; Hoko (Kuroda, 1941, p. 111; y); Karenko; Lasy- oto; Garanbi; Hoko; Ryukyu-syo (Kuroda, 1941, p. 111; w). RYUKYU ISLANDS: Kikaiga Shima (ANSP; y); Bolo reef, NW Nakagami Gun, Okinawa (USNM; y); Okuma, Kuni- gami-Gun; Odomari, Okinawa (both USNM; w); Yomitan reef, Okinawa (AMNH; yw). JAPAN: Oshima, Osumi (USNM; yw); Hachijo Id., off Honshu (ANSP; w); Tosa, Shikoku (ANSP; y); Kagoshima, Kyushu (ANSP; w). MARI- ANAS: Lagunan Tanapaa, Saipan; Agat Bay, Guam (both ANSP; y); Apra Harbour, Guam (AMNH; y); Tinian Id. (FMNH; yw); several localities on Guam Id. (USNM; w); Saipan (AMNH; w). PALAU ISLANDS: Angupelu Id., SE Koror Id.; reef N. of Gorokottan Id., S. side W. Passage, Babelthuap Id. (both ANSP; y); S. E. of Auropushekaru Id., Malakal Harbour, Koror Id. (ANSP; yw); Angaur, Peleliu Id. (FMNH; w). CAROLINE ISLANDS: N. of Kayangel Id.; S. of Garakayo Id.; Ponape; Yap (all ANSP; y); Ulithi Atoll; Satawal Atoll; Manini, Kapingamarangi (all USNM; y); S.E. Rattakadokoru Id. (ANSP; w). MARSHALL ISLANDS: Eni- wetok Atoll; Kwajalein Atoll (both AMNH; y); Bikini Atoll; Eniwetok Atoll; Rongelap Atoll; Rongerik Atoll; Kwajalein MicoBAR |: MaLoy 13 ve et ot © of mazamae's, TRICINUS RICINUS 160° 180° 160° igo T T 7 90° 120° 100° r HEBRID earn | | | @ YELLOW-SPOTTED FORM © 80TH FORMS TEM Ap O WHITE FORM © UTERATURE RECORD l — | 60" ie ey = see T25* Plate 17. Geographical distribution of Drupa (Drupa) ricinus ricinus (Linnaeus) in the Indo-Pacific, and its subspecies D. (D.) ricinus hadari Emerson and Cernohorsky, in the Red Sea. [14-826] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 23 Atoll (all ANSP; w). WAKE ISLAND: (AMNH, yw; ANSP, y). NEW GUINEA: NE Noekori Id.; Manokwari; SW Biak dock, Biak Id. (all ANSP; y); Boensaki Id., off Sowek, Soepi- ori Id., Schouten Ids. (ANSP; yw); NE Mioes Woendi, Pad- aido Ids. (ANSP; yw); Samarai, Papua (AMNH; yw); Huon Gulf (USNM; w). AUSTRALIA: Queensland: Capricorn Ids.; Hardy reef, Great Barrier reef (both AMNH; y); Birds Id.; Green Id., Cairns (both USNM; y); Magnetic Id.; Lady Elliot Id. (both AIM; y); New South Wales: (Iredale & McMichael, 1962, p. 74); West Australia: Vlaming Head (FMNH; y); Houtman, Abrolhos Archipelago (Wilson & Gillett, 1971, p. 92). NEW BRITAIN: Rabaul Harbour (ANSP; y). NEW IRELAND: Kavieng (AMNH; y). SOLOMON ISLANDS: Kieta, Bougainville; Lutee, Choiseul Id.; Ataa, N. Malaita Id.; Tulagi Id., Santa Ana Id. (all AMNH; y); Tikopia Id. (AIM; w); Lunga, Guadalcanal (AMNH; yw). NEW HEB- RIDES: Lamap, Malekula Id. (ANSP; w); Bushmens Bay, E. Malekula Id. (AIM; w); Pango Point, Efate Id. (AIM; w); Tauna Id. (DM; w). NEW CALEDONIA: SE Dumbea Pass, off Noumea; Baie Quemo, Noumea; Koe, Touho (all ANSP; y). LOYALTY ISLANDS: Lifu (USNM; y). GILBERT IS- LANDS: Abemama (USNM; w). ELLICE ISLANDS: Funa- futi lagoon (USNM; AIM; w). WALLIS & FUTUNA IS- LANDS: Faioa; Uvea; Nukuhifala, all Wallis Id. (USNM; y); Nukuhifala, Wallis Id. (USNM; w); Anse de Sigave, Futuna Id. (USNM; yw); Mua, Alofi, Hoorn Id. (USNM; y). FIJI ISLANDS: Koro Bay reef, Vanua Levu (AMNH; y); Suva Harbour, S. Viti Levu (USNM; y); Korolevu, S. Viti Levu; Yasawa reef (both AMNH; w); Namagumagua village, S. Viti Levu (WOC coll.; w); Rat Tail Passage, Suva reef, S. Viti Levu; Mamanuca_ group; Caboni beach; Manava_ Id.; Nananu-i-Ra Id.; Viti Levu Bay, all Viti Levu (all WOC coll.; yw). TONGA ISLANDS: Ha’ateiho reef, Tongatapu; E. coast Tongatapu Id. (both USNM; y); Haakoma, Tongatapu (MCZ; w); Hufagalupe, Tongatapu (MCZ; yw) Niuafo’ou Id. (USNM; w). NIUE ISLAND: near Malatu (USNM; y); Oneone reef (AMNH; yw); Alofi (USNM; yw). SAMOA ISLANDS: Vailele Bay, Upolu; Tafuna, Tutuila (both ANSP; y); Fagaitua Bay, Tutuila; Ofu (both MCZ, w); Asau Harbor, Savaii (USNM; yw); Apia, Upolu (AMNH; yw); Tau Id., Manua group (MCZ; yw). TOKELAU ISLANDS: W. side Tukuo, Manihiki Atoll (ANSP; w); Swains Id. (USNM; w). PHOENIX ISLANDS: Canton Id.; Enderbury Id. (both USNM; w). HOWLAND IS- LAND: (USNM; w). COOK ISLANDS: Bird’s Id., Tom’s Id.; North Id., all Palmerston Atoll (all USNM; w); Koromiri Id., SE Rarotonga (ANSP; y); several localities on Aitutaki; Mauke; Rarotonga; Mangaia (all MCZ; USNM; w); Motu Akaiami, Aitutaki (USNM; yw). AUSTRAL ISLANDS: Rurutu; Tubuai; Raivavae (all USNM; w). SOCIETY IS- LANDS: Mopelia Id. (ANSP; y); many localities on Bora Bora; Huahine; Raiatea; Moorea; Tahiti (USNM; w); Arue, Tahiti; Taone, Tahiti; Huahine (all USNM; yw). TUAMOTU ISLANDS: Napuka (AMNH; w); Tacume; Clermont Tenere; Takaroa (all ANSP; w); Rangiroa (AMNH; yw); Mangareva (USNM; w); Raroia (ANSP; yw). PITCAIRN ISLAND: (AMNH; USNM; w). MARQUESAS ISLANDS: Taiohae, Nukuhiva (AMNH; y); Hiva Oa Id. (ANSP; w); Ua Huka Id. (AMNH; ANSP; w). LINE ISLANDS: Christmas Id.; Flint Id.; Palmyra Id. (all ANSP; w); Palmyra Id. (MCZ; y); Jarvis Id. (AMNH; y). JOHNSTON ISLAND: (USNM; w). HAWAI- IAN ISLANDS: Honolulu Harbour (ANSP; y); Kure Id.; Midway Id.; Laysan Id.; Tern Id.; French Frigate Shoal; Kauai Id.; Oahu Id.; Molokai Id.; Maui Id.; Hawaii Id. (all AMNH,; ANSP; USNM; w); Niihau Id.; Oahu Id. (both AIM; w); Lihue, Kauai (AMNH; yw). CLIPPERTON ISLAND: (AMNH,; ANSP; w); (Hertlein & Allison, 1960, p. 15; yw). GALAPAGOS ISLANDS: (Hertlein, 1960, p. 8). Fossil records— ZANZIBAR: Pleistocene: Base of well, vil- lage W. of Makunduchi (Cox, 1927, p. 90, pl. 18, figs. 12a, b and Stockley, 1928, p. 79). MOZAMBIQUE: Pleistocene: Chidenguel, Inhambane district (Cox, 1939, p. 90). HAWAI- IAN ISLANDS: Pleistocene: Oahu (Mansfield in Stearns and Vaksvik, 1935, p. 167; Kosuge, 1969, p. 786, pl. 5, fig. 90). Drupa ricinus new subspecies hadari Emerson and Cernohorsky (PI. 2, figs. 9, 10; Pl. 18) Range—Red Sea only. Remarks—Distinguished from the nominate subspecies Drupa_ ricinus (Linnaeus) by _ its larger size, heavier shell and more strongly de- veloped parietal shield. Habitat—Coral reefs and rocks, on intertidal reef-flats. Description—Juveniles typical of those of both apertural color forms of the nominate subspecies, with spinose nodules dark brown. Nodules on the body whorl or mature specimens white, with brown nodules on spire only, or body whorl ax- ially streaked with brown except on the last two or three rows of spines. Aperture in mature spec- imens white with fully developed parietal shield completely concealing nodular coloration of the parietal area. The side-cusps of the rachidian of the radula are bifid and are followed by 2 weak, short lat- eral denticles, excluding the slightly longer end- cusps. Measurements (mm)—(including spines) length width 38.0 34.2 Eilat, Gulf of Aqaba (paratype AMNH no. 112617b) 37.2 38.7 Eilat, Gulf of Aqaba (holotype AMNH no. 166928) 34.2 34.1 Eilat, Gulf of Aqaba (paratype AMNH no. 112617a) 32.0 32.0 Eilat, Gulf of Aqaba (paratype AMNH no. 112617c) 28.0 29.0 Eilat, Gulf of Aqaba (paratype DMNH no. 51119) Synonymy— 1862 Ricinula albolabris Blainville, Ktister (pars), Syst. Conchylien-Cabinet, ed. 2, Abt. IE, Ricinula, pl. 2, figs. 1, 2 (Red Sea) [non Purpura albolabris Blain- ville, 1832]. 1941 Drupa (Drupa) ricinus (Linné), Wenz, Handb. Pali- ozoologie, vol. 6, pt. 5, p. 1114, fig. 3165 (Red Sea) {non Murex ricinus Linnaeus, 1758]. Pe pa Se Can aaa : nN “hy | J V/ Plate 18. Radula of Drupa (Drupa) ricinus hadari Emerson and Cernohorsky. Half a transverse row; Eilat, Gulf of Aqaba, Israel. [14-827] 24 = Drupa Types—The holotype, AMNH No. 166928, and 8 paratypes AMNH No. 112617, have been de- posited in the American Museum of Natural His- tory, New York. Other paratypes are in the Dela- ware Museum of Natural History, Greenville, The Auckland Institute and Museum and the Tel Aviv University, Israel. The type locality is Eilat, Gulf of Aqaba, Israel. Nomenclature—The subspecies is named in honor of the late Aryeh Hadar, who kindly sub- mitted specimens from the Gulf of Aqaba for study. Records—RED SEA: Sharem, Gulf of Suez (DMNH): Eilat, Gulf of Aqaba (AMNH; DMNH; AIM; w); Ras Banas, Egypt (USNM; w); Quseir, Egypt (LACMNH; y); Mualla, Abu Zabad, Gulf of Aqaba (Rees and Stuckley, 1952, p. 196); Suakin, Sudan (Sturany, 1905, p. 141); Jiddah, Saudi Arabia (AMNHII; yw); Pointe du Requin, [le Abulat (Franc, 1956, p. 37). Drupa elegans (Broderip and Sowerby, 1829) (PI. 2, fig. 12; Pl. 19, 20) Range—From Wake Island to the Tuamotu Is- lands. Remarks—This species is similar to Drupa ri- cinus (Linnaeus), but differs in apertural color- ation, the lack of tubercle coloring and in being smaller in size. This uncommon species occurs sympatrically throughout its range with D. ri- cinus (Linnaeus). It may prove eventually to be another, moderately rare color phase of that variable species. Although an occurrence of D. elegans in the Society Islands is probable, the cited record requires confirmation; specimens from that locality in the Academy of Natural Sciences of Philadelphia and the National Mu- seum of Natural History, Washington, lack exact locality data. Habitat—Only five specimens were taken by the operation “Crossroads” in 1946 and the biologi- cal re-survey in 1947 of the Marshall Islands. These were found living in association with the extremely common _ yellow-spotted and_ white color phases of Drupa ricinus. On Niue Island, the species is also sympatric with D. ricinus and has been collected on reefs under cliff overhangs (D. C. Johnson, personal commun.). Description—Shell 13 to 25 mm (about % to 1 inch) in length, sub-ovate, spire short; body whorl spirally ribbed with five rows of siphonous Emerson and Cernohorsky Muricidae Limam Plate 19. Radula of Drupa (Drupa) elegans (Broderip and Sowerby). Half a transverse row; Vailoa, Alofi Bay, Niue Id. Plate 20. Drupa (Drupa) elegans (Broderip and Sowerby). Figs. 1, 2. Lectotype from unknown locality (B.M. (N.H.), 20.3 x 22.4 mm) [photo courtesy J. Taylor, B. M. (NH)]. Figs. 3, 4. Specimen from Vailoa, Alofi Bay, Niue Id. (WOC coll.; 19.0 x 20.7 mm). tubercles. Intersticial surface minutely scaled. Parietal shield moderately enameled, reflected over body wall to form a heavy callus; lower por- tion of columella 4 plaited. Axial fold strongly developed. Aperture very narrow, dentition of [14-828] May 30, 1973 outer lip as in Drupa ricinus (Linnaeus). Color of the exterior and aperture white; aperture encir- cled by a continuous dark riddish brown line in mature specimens; in juvenile specimens, the line is disrupted and restricted to the columellar lip and near the anal siphonal canal. Radular ribbon very small, side-cusps of the rachidian bifid, followed by 3. slender and deeply rooted lateral denticles, exclusive of the short end-cusps. Measurements (mm)—(including spines; all specimens with a mature lip) length width 25.0 25.0 Wake Island 20.3 22.3 Lectotype of elegans B. and S. 19.0 20.7 Niue Id. 18.5 17.5 Lord Hood Id. (S. Marutea Id.) 13.6 11.8 Lord Hood Id. (S. Marutea Id.) Synonymy— 1829 Ricinula elegans Broderip & Sowerby, Zoological Jour- nal, London, vol. 4, p. 376 (no locality given); 1839 Gray, Zool. Capt. Beechey’s voyage, p. 155, pl. 36, fig. 4; 1844 Deshayes & Milne-Edwards, Hist. nat. anim. s. verttbres, ed. 2, vol. 10, p..52; 1846 Reeve, Conchologia Iconica, vol. 3, pl. 1, fig. 1 (Lord Hood Id. = South Marutea Id.); 1933 Dautzenberg & Bouge, Journal de Conchyliologie, vol. 77, p. 238. 1853 Pentadactylus elegans Broderip, H. & A. Adams, Gen- era Recent Mollusca, vol. 1, pp. 129, 130; vol. 3, pl. 13, figs. 6a, b (operculum only). 1880 Ricinula ricinus var. elegans Broderip, Tryon, Manual of Conchology, vol. 2, p. 184, pl. 56, fig. 193. 1913 Drupa ricinus var. elegans Broderip & Sowerby, Hed- ley, Nautilus, vol. 27, no. 7, p. 80. 1957 Drupa elegans Broderip & Sowerby, Kaicher, Indo- Pacific Sea Shells, (Muricacea, Buccinacea), pl. 3, fig. 18. Types—Three syntypes of Drupa elegans are in the British Museum (Nat. Hist.), and the 20.3mm long syntype (PI. 20, figs. 1, 2) is here selected INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa = 25 as the lectotype. The species was described from shells in the Museum of the Zoological Society brought home by Lieutenant Belcher who sailed under Captain Beechey in the “Blossom” on its voyage to the Bering Straits and the Pacific (1825-1828). No type locality was given, but Cuming collected it from Lord Hood Island (= S. Marutea Id.), and his specimen was figured by Reeve (1846). Lord Hood Island (= S. Marutea Id.) is here designated as the type locality. Records—WAKE ISLAND: (BPBM). MARSHALL IS- LANDS: Namu Id., N. W. end of Bikini Id.; Eniman Id., Bikini Atoll; Eniwetok Atoll (all USNM). LINE ISLANDS: Caroline Id. Flint Id. (both ANSP). NIUE ISLAND: (AMNH); Vailoa, Alofi Bay (D. C. Johnson coll.; WOC coll.); Avatele (S. Herriot coll.). COOK ISLANDS: Akamaru Id., Manihiki Atoll (ANSP). SOCIETY ISLANDS: (ANSP; USNM). TU- AMOTU ISLANDS: Lord Hood Id. (= S. Marutea_ Id.) (AMNH; BMNH); Vahitahi; Nengonengo Id. (both USNM); Anaa Id. (MCZ); Napuka; Taenga; Fakahina (Dautzenberg & Bouge, 1933, p. 238). 140° 160° 180° 160° 140° 120° CAROLINE wove TELEGANS 6.4 —_| = NEW ZEALAND Sp | 1 160° 180° Plate 21. Geographical distribution of Drupa (Drupa) ele- gans (Broderip and Sowerby). Open circles are literature records. [14-829] 26 Drupa Emerson and Cernohorsky Muricidae ps ae ce en ie Ae [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence. } [14-830] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa = 27 Subgenus Ricinella Schumacher, 1817 Type: Drupa rubusidaeus Réding, 1798 Shell sub-ovate, solid, spire rather short; body whorl large, whorls spirally ribbed, sculptured with siphonous spiniform tubercles; interstices filled with fine scale-like plates forming raised threads; threads running spirally in close parallel association; columella with an axial fold, outer lip dentate with teeth singularly arranged, some- times in close association, not compound; oper- culum subelliptical, not linear; radula typical for the genus. This group includes besides the type species, Ricinula speciosa Dunker, 1867, R. clathrata clathrata Lamarck, 1816, and R. clathrata miti- cula Lamarck, 1822. On the basis of radular characters, the group shows a close _relation- ship with Drupa sensu stricto. The shells on the other hand resemble some of the thaids. The members of this group have the labial teeth de- veloped as small, pearl-like teeth which may be closely crowded together as in Drupa clathrata or D. speciosa, but are never actually compound. The interpretation of the identity of the type species of Ricinella, i.e. R. purpurata Schu- macher, rests on the elucidation of the cited illus- tration of Favanne (1784, pl. 24, fig. 2). This fig- ure resembles D. rubusidaeus Riding more so than D. clathrata (Lamarck). Favanne’s descrip- tion of the color of the aperture as being deep and vidid lilac or purple, excludes the species D. clathrata from consideration. Synonymy— 1817 Ricinella Schumacher, Essai nouv. systéme, pp. 72, 240. Type-species by subsequent designation, Iredale, 1937: Ricinella| purpurata Schumacher, 1817 [= Drupa rubusidaeus Réding, 1798]. | pa Plate 22. Radula of Drupa (Ricinella) rubusidaeus Réding. Half a transverse row; Olawala, Maui, Hawaiian Ids. Drupa rubusidaeus Réding, 1798 (Pl. 2, figs. 13-15; Pls. 22, 23) Range—From the Red Sea and East Africa throughout the Indo-Pacific to Hawaii and the Tuamotu Islands. Remarks—This solid species could only be con- fused with the much smaller and rarer Drupa speciosa (Dunker). D. speciosa is a higher-spired shell with no yellow coloration in the aperture; the pink coloring of the aperture is more pro- nounced and the denticles of the outer lip are more closely set and almost grouped. Habitat-—On algae matted reef-flats, under rocks and in tide-pools, generally on the seaward half of the reef-flat; from low tide to a depth of 10 fathoms or more, on rock, sand or coral. In the Hawaiian Islands the species has been col- lected attached to stony coral at a depth of 50 to 60 feet (C. S. Weaver, leg.). Description—Shell 20 to 55 mm (% to 2% inches) in length, solid, ovately globose, slightly ventri- cose, spire short, acuminated, body whorl large and with five rows of strong, sub-spiniform, siphonous tubercles. Sculpture of fine, scale like Plate 23. Drupa (Ricinella) rubusidaeus Réding. Fig. 1. Lectotype figure of D. (R.) rubusidaeus Rodding, from Knorr, 1768, pt. 6, pl. 24, fig. 7. Fig. 2. Holotype of Ricinula reeveana Crosse, from Nuku- hiva, Marquesa Ids. (B.M. (NH); 48.2 x 38.8 mm—im- mature specimen). [14-841] 28 Ricinella plates forming ridges which run spirally in par- allel association between the rows of tubercles. Aperture sub-elliptical, outer lip in adult stage dentate with 7-12 small pearl-like teeth; margin crenulated, apertural area enameled; enameled area extending over a large portion of the body whorl adjacent to the aperture. Inner lip cal- lused with 3 or 4 plications. Siphonal canal short, deep, recurved; sutural canal conspicuous, always open. Color white-yellowish externally, margin of the aperture yellow, interior of the aperture rich pink. The side-cusps of the rachidians of the radula are bifid to trifid and are followed by 2-3 lateral denticles, exclusive of the stronger side-cusps. Measurements (mm)—(including — spines; all specimens with a mature lip) length width 53.8 48.5 Oahu, Hawaiian Ids. 48.2 38.8 Holotype of reeveana Crosse 41.6 38.9 Luzon Id., Philippines 34.0 28.7 Niue Id., Polynesia 28.8 25.0 Koror Id., Palau Ids. Synonymy— 1742 Gualtieri, Ind. test. Conchyliorum, pl. 28, fig. R (poor). Regenfuss, Aus. Schnecken, Muscheln u.a.Schaal- thiere, (1), pl. 3, fig. 32. Knorr, Verg. Augen U. Gemiiths, pt. 6, pl. 24, fig. 7 (very good). 1777 “Murex hericinus” Martini, Syst. Conchylien-Cabinet, vol. 3, p. 283, pl. 101, figs. 974, 975 (poor). [non- binomial]. Favanne, La Conchyliologie d’Argenville], pl. 24, fig. A2 (poor). Favanne, Catalogue systematique et raisonné ee ates p. 145 (description). 1791 Murex nodus Gmelin, Systema Naturae, ed. 13, p. 3537 (refers to Knorr, op. cit., with a query ) [no locality given] (non Linnaeus, 1758). 1798 Drupa rubusidaeus Roéding, Museum Boltenianum, p. 55 (refers to Martini, op. cit., and Knorr, op. cit.) [no locality given]; 1913 Hedley, Nautilus, vol. 27, no. 7, p. 80; 1966 Weaver, Hawaiian Shell News, vol. 14, no. 14, p. 2, textfigs. 1, 2; 1969 Cernohorsky, Veliger, vol. 11, no. 4, p. 301, pl. 47, figs. 10, 10a; 1971 Wilson & Gillet, Australian Shells, p. 92, pl. 61, figs. 1, la. , 1798 Drupa fragum hoding, Museum Boltenianum, p. 55 (refers to Murex nodus Gmelin, 1791, sp. 42) [no locality given]. 1807 Mancinella hystrix Link, Beschr. Nat.-Samml. Univ. Rostock, 3 Abth., p. 115 (refers to Martini, op. cit.) [non Murex hystrix Linnaeus, 1758]. 1817 Ricinella purpurata Schumacher, Essai nouv. systéme, p. 240 (refers to Favanne, op. cit.) [no locality given]. 1817 Murex hystrix Linnaeus, Dillwyn, Desc. cat. Rec. shells, vol. 2, p. 706 (refers to Martini, Gualtieri, Regenfuss etc.) [East Indian Seas; coasts of the Friendly Islands = Tonga Ids.]; 1825 Wood, Index Testaceologicus, p. 124, pl. 26, fig. 50a (non M. hystrix Linnaeus, 1758). 1758 1768 1780 ed. 3. [of 1784 Emerson and Cernohorsky Muricidae 1822 Purpura hystrix Lamarck, Hist. nat. anim. s. vertebres, vol. 7, p. 247 (refers to Knorr, Regenfuss, Martini and Gualtieri, op. cit.); 1835 Kiener, Spéc. gén. icon. coq. viv., vol. 8, p. 13, pl. 2, figs. 4, 4a, b; 1846 Reeve, Conchologia Iconica, vol. 3, pl. 3, fig. 13 (non Murex hystrix Linnaeus, 1758). 1825 Murex hippocastanum Wood, Index Testaceologicus, p. 124, pl. 26, fig. 53a (non Linnaeus, 1758). 1832 Purpura spathulifera Blainville, Nouv. Ann. Mus. (Hist. Nat. Paris, ser. 3, vol. 1, p. 212, pl. 9, fig. 8 (no locality given). 1833 Purpura histrix (sic) Lamarck, Quoy & Gaimard, Voy- age L’Astrolabe, vol. 2, p. 575, pl. 39, figs. 14-16 (animal and operculum). 1853 Pentadactylus hystrix H. & A. Adams, Gen. Rec. Mol- lusca, vol. 1, p. 130; 1875 Troschel, Gebiss d. Schnecken, vol. 2, p. 134, pl. 13, fig. 4 (radula). 1862 Ricinula reeveana Crosse, Journal de Conchyliologie, vol. 10, p. 47, pl. 1, fig. 3 (Nouhiva = Nukuhiva, Mar- quesas Ids.) [immature specimen] (non Ricinula reeveana C. B. Adams, 1852). 1880 Ricinula hystrix Linné, Tryon, Manual Conchology, vol. 2, p. 183, pl. 56, tig. 195; 1933 Dautzenberg & Bouge, Journal de Conchyliologie, vol. 77, p. 239. 1880 Ricinula hystrix var. reeveana Crosse, Tryon, ibid., vol. 2, p. 183, pl. 56, fig. 196; 1933 Dautzenberg & Bouge, ibid., vol. 77, p. 239 (non C. B. Adams, 1852). 1911 Sistrum hystrix Linné, Schepman, Siboga-Expeditie, vol. 49d, p. 354 (non Murex hystrix Linnaeus, 1758). 1913 Drupa rubusidaeus Bolten, Hedley (pars), The Nau- tilus, vol. 27, no. 7, p. 79. 1936 Drupa spathulifera (Blainville), Hirase, Coll. Jap. shells, ed. 5, p. 79, pl. 110, fig. 9; 1954 Kira, Col. illust. shells Japan, p. 47, pl. 23, fig. 4; 1967 Habe & Ko- suge, Stand. book Jap. shells color, vol. 3, p. 70, pl. 27, fig. 24; 1968 Taylor, Phil. Trans. Roy. Soc. London, ser. B, vol. 254, p. 201. 1938 Drupa (Drupa) hystrix (Linné), Adam & Leloup, Mém. Mus. Roy. d’Hist. Nat. Belgique, vol. 2, fasc. 19, p. 163. 1957 Drupa rubuscaesia Kaicher, Indo-Pacific Sea Shells (Muricacea, Buccinacea), pl. 3, fig. 15 (non D. rubus- caesius Réding, 1798). 1961 Drupa rubuscaesium Rodding, Rippingale & McMichael, Queensland and Gt. Barrier reef Shells, p. 102, pl. 13, fig. 3 (non D. rubuscaesius Roding, 1798). 1962 Drupa rubuscaesius (R6éding), Kira, Shells West. Pa- cific in color, vol. 1, p. 62, pl. 24, fig. 4; 1965 Ara- kawa, Venus: Jap. Journ. Malacology, vol. 24, no. 2, p. 115, pl. 13, figs. 5, 6 (radula) [non Réding, 1798]. 1965 Drupa speciosa (Dunker), Wu, Bull. Inst. Zool. Acad. Sinica, vol. 4, p. 98, textfig. 30 (radula) [non Ricin- ula speciosa Dunker, 1867]. 1967 Drupa rubusidaea Réding, Orr Maes, Proc. Acad. Nat. Sci. Philadelphia, vol. 119, no. 4, p. 129. T ypes—Réding’s type specimen of Drupa rubu- sidaeus is lost, and we therefore select the spec- imen figured by Knorr on plate 24, fig. 7, as the lectotype of the species (Pl. 23, fig. 1), and designate Davao Bay, Mindanao, Philippines, as the type locality. The type specimen of Purpura spathulifera Blainville, is probably in the Mu- séum National d’Histoire Naturelle, Paris, and the holotype of Ricinula reeveana Crosse, is in the British Museum (Nat. Hist.) [Pl. 23, fig. 2]. The type specimen of Ricinella purpurata Schu- [14-842] May 30, 1973 o9° INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa_ 29 20°|— ees: or RUBUSIDAEUS] ‘y CHAOS s.? Plate 24. Geographical distribution of Drupa (Ricinella) rubu- sidaeus Réding. Open circles are literature records. macher, could not be located in the Zoological Museum, Copenhagen (J. Knudsen, in litt.). Nomenclature—This species was at first mas- querading under the names Mancinella hystrix Link, Murex hystrix Dillwyn and Purpura hystrix Lamarck. As previously discussed, this is not Murex hystrix Linnaeus, 1758. Since Réding, 1798, cites 2 of the 4 references which Lamarck considered synonyms of Purpura hystrix, there can be no doubt regarding the identity of his species. Réding’s citation to Knorr’s excellent figure (designated as lectotype) of the present species, as well as Martini’s figures, serves to establish the identity of D. rubusidaeus. More recently Hedley (1913) confused Purpura spathulifera Blainville, a species which he synon- ymized with Drupa rubuscaesius Roding, Ricin- ula clathrata Lamarck, and R. speciosa Dunker, but which is conspecific with D. rubusidaeus. The specimen and its figured radula cited by Wu (1965a) as “Drupa speciosa (Dunker)” originated from Wan-li-tong, Taiwan, and is actually D. rubusidaeus Réding (Wu in litt.). Records—RED SEA: (AMNH). EAST AFRICA: Gesira, Somalia (ANSP); Diani Beach, Kenya (Heinicke, 1970, p. 7); Kendwa Id., 4 mi. ESE of Dar-es-Salaam; Sinda Id., 15 mi. SSE of Dar-es-Salaam; N.W. of Magogani; Pangavini Id., 10 mi NNE of Dar-es-Salaam, all Tanzania (all MCZ); Mozam- bique (USNM). SEYCHELLES ISLANDS: Beau Vallon Beach, Mahe; Cousin Id. (both ANSP). MADAGASCAR: Nosy N’Tangam, W. Nossi Bé; Grande Recife, W. of airport, Tulear; Ambodifototra (all MCZ). REUNION ISLAND: (ANSP). MALDIVE ISLANDS: Miladummadula Atoll; Tila- dummati Atoll; N. Malé Atoll (all ANSP). LACCADIVE ISLANDS: (Hornell, 1922, p. 217). CEYLON: (Langdon, 1875, p. 72). VIETNAM: Cam Ranh Bay (ANSP). COCOS- KEELING ISLANDS: Klapetuju, West Id. (USNM); S. end of Home Id.; N. end of Horsburgh Id.; S. end of Direction Id. (all ANSP). CHRISTMAS ISLAND: (Tomlin, 1935, p. 79). INDONESIA: Mandi Darrah Id., N. Borneo (AMNH; ANSP); Pulau Bai, Batu group, off Sumatra; Pulau Penju, S. Sumatra (both USNM); Bali Id.; Wasi, Ambon Id. (both FMNH); Banda Id.; Soengai Manoembaii, Hes Aroe (Adam & Leloup, 1938, p. 164); Tjilaoet, Java (Altena, 1945, p. 146). PHILIPPINES: Iba, Zambales, Luzon Id.; Boron- gan village, E. Samar Id.; Cuyo Id., Palawan group (all ANSP); Tilig reef, Lubang Id.; Calapan, Mindoro: Linga- yan Gulf; Davao Bay, Mindanao (all AMNH); Nogas Point, Panay Id.; Cabra Id.; Silino Id.; Point Matangal, Basilan, Sulu (all USNM). FORMOSA (Taiwan): Wan-li-tong (AMNH); Karenko; Botan-wan; Ryuku-syo; Hoko (Kuroda, 1941, p. Ill). RYUKYU ISLANDS: Yomitan’ reef, Okinawa (AMNH); Okuma, Kunigami-Gun; Bolo reef, N.W. Naka- gami-Gun (both USNM); Ishigake (MCZ); Miyako (FMNH). JAPAN: Hachijo Id., 275 mi. S. of Tokyo; Tosa, Shikoku; Kagoshima, Kyushu (all ANSP); Ominato Ise (AIM); Oshima, Osumi (Powell coll.). MARIANAS: Saipan (AMNH); Agana Bay, Guam Id. (ANSP). PALAU ISLANDS: Babelthuap Id.; Angupelu Id., S.E. of Koror; Malakal Harbour, Koror; Helen Channel (all ANSP). CAROLINE ISLANDS: Kayangel; Ngargersiul; S.W. of Rattakadokoru; S. of Garakayo; Ulithi; Ella; Elangalap Id., Ifaluk Atoll; Tirakaume; Ringutoru; Kapingamarangi (all ANSP); Ponape reef (AMNH). WAKE ISLAND: (BPBM). MARSHALL ISLANDS: Eniwetok Atoll, Bikini Atoll; Rongerik Atoll (all USNM); Kwajalein Atoll (Dietrich & Morris, 1953, p. 15). NEW GUINEA: Biak Id., Schouten Ids. (USNM); 1 mi. NE of Mioes Woendi, Padaido Ids. (Powell coll.); Wewak (ANSP). NEW BRITAIN: Rabaul Harbour (AMNH). AUSTRALIA: Queensland: Herald Cay, Coral Sea; Holmes reef, Coral Sea; off Cairns; Watt reef, off Townsville (all AMS); Green Id., off Cairns (AIM); several islands of the Capricorn group (AMS; AMNH). SOLOMON ISLANDS: Choiseul Bay, Choiseul Id.; Bougainville Id.; Bellona Id. (all AMNH); Fiu, Malaita Id. (Powell coll.). NEW HEBRIDES: Efate Id. (Colardeau coll.); Espiritu) Santo (Solem, 1959, p. 262). NEW CALEDONIA: Touho (AMNH). GILBERT ISLANDS: Abaiang (MCZ); Kingsmill Ids. (AMNH). ELLICE ISLANDS: Funafuti lagoon (AIM). FIJI ISLANDS: (AMNH,; ANSP); Rat Tail Passage, Suva reef, S. Viti Levu (WOC coll.); Mamanuca group (Jennings coll.). NIUE ISLAND: (DM); Tuapa reef (AMNH; WOC coll.). SAMOA ISLANDS: reef at Satalo Id., Upolu Id.; E. side of Wailele Bay, Upolu Id. (both ANSP); Nuuli, Tutuila Id. (MCZ). COOK ISLANDS: Mauke; Aitutaki (both USNM); off Aroa Creek, S.W. Rarotonga; Koromiri Id., S.W. Raro- tonga (both MCZ); Mangaia (DM). SOCIETY ISLANDS: Several localities on Tahiti and Moorea (USNM). TUA- MOTU ISLANDS: Anaa Id. (AMNH; ANSP; MCZ); Amanu (Coututirer, 1907, p. 143). MARQUESAS ISLANDS: (MCZ); Nukuhiva (BMNH). LINE ISLANDS: Fanning Id. (DMNH). HAWAIIAN ISLANDS: Midway Id.; off Waikiki, Oahu (both [14-843] 30 Ricinella Emerson and Cernohorsky Muricidae AMNH); Makaha Point, $.W. Oahu (MCZ); Olawala, Maui (AMNH). Fossil records—KENYA: Pleistocene: raised reef, S.E. of Mombassa Id., N. of Ras Serani (Cox, 1930, p. 145). HA- WAITAN ISLANDS: Oahu Id. (Kosuge, 1969, p. 786, pl. 5, fig. 94). Drupa speciosa (Dunker, 1867) (Pl. 2, figs. 21, 22; Pl. 25) Range—Tuamotu and Pitcairn Islands. Remarks—The shell of this Polynesian species is smaller and higher-spired than D. rubusidaeus Réding, with a deeper pink to mauve aperture which lacks the yellow coloration found in the aperture of the larger species. Specimens labelled “Fiji Islands” are obviously based on erroneous locality indications as the species does not occur there. The record from Rarotonga, Cook Islands (ANSP) is also suspect, and has not been con- firmed by recently collected, well-documented specimens. Habitat—Unknown, but probably found on coral reefs at low tide. Description—Shell 20 to 29 mm (% to 14 inches) in length, solid, ovate, globose; spire moderately elevated, acuminate; body whorl axially ribbed with 9 ribs, crossed by 5 transverse rows of short, strong, spinose tubercles. Interstices be- tween tubercles striated with parallel rows of small scales. Aperture subelliptical, columella posteriorly excavated, outer lip dentate with 4-7 evenly spaced small white teeth; margin crenu- lated. Inner lip with 3 strong lower plications and 1 subobsolete upper plication. Apertural area enameled, enameled area of the inner lip extend- ing over a portion of the body whorl. Anterior siphonal canal short, deep; posterior siphonal canal open. Color creamy-white externally, aper- ture a deep mauve pink. Radula unknown. The radular dentition figured by Wu (1965a) for this species was based on a specimen of D. rubusidaeus from Taiwan (Wu, in litt.). Measurements (mm)—(including spines; all specimens with a mature lip) length width 27.0 21.5 Tuamotu Ids. 26.9 23.2 Lectotype of speciosa Dunker 26.4 20.8 Raroia, Tuamotu Ids. 25.4 21.7 Vahitahi, Tuamotu Ids. 22.8 18.9 Vahitahi, Tuamotu Ids. 21.9 16.3 Vahitahi, Tuamotu Ids. 20.1 14.8 Tuamotu Ids. Synonymy— 1846 Ricinula clathrata Lamarck, var. B., Reeve, Conchol- ogia Iconica, vol. 3, pl. 2, fig. 9a (Anaa Id.) [non R. clathrata Lamarck, 1816]. 1867 Ricinula speciosa Dunker, Novitates Conchologicae, Abt. I, pts. 11/12, p. 100, pl. 33, figs. 7, 8; 1878 Dunker, Addenda & Corrigenda, p. 139 (Philippine Ids. = error) [non Purpura speciosa Valenciennes, 1832]. 1880 Ricinula hystrix var. speciosa Dunker, Tryon, Manual Conchology, vol. 2, p. 183, pl. 56, fig. 194; 1933 Dautzenberg & Bouge, Journal de Conchyliologie, vol. 77, p. 239. 1969 Drupa speciosa Dunker, Cernohorsky, Veliger, vol. 11, no. 4, pp. 301, 302. Types—Dunker described the species from 4 specimens in Hugh Cuming’s collection which is now in the British Museum (Nat. Hist.). Three of the syntypes are immature examples with weak denticles on the outer lip and a superficial col- umellar callus. Only one specimen is reasonably mature, and this specimen is here selected as the lectotype of Ricinula speciosa Dunker (P1 25, fig. 1). Even though the selected lectotype is without Plate 25. Drupa (Ricinella) speciosa (Dunker). Fig. 1. Lectotype (B.M. (NH); 26.9 x 23.2 mm). Fig. 2. Syntype; immature (B.M. (NH); 26.9 x 21.9 mm) {photos courtesy K. Way, B.M. (NH) ]. Fig. 3. Specimen from Vahitahi, Tuamotu Ids. (USNM 613343; 22.8 x 19.0 mm). [14-844] May 30, 1973 question the Drupa speciosa of authors, the spec- imen does not compare too well with Dunker’s original type-figure. Either the artist exercised his own imagination when depicting the spec- imen, or the originally illustrated example has gone astray. Stability of the taxon Drupa speciosa, however, is best served by the selection of the most mature specimen from among the 4 syn- types, which all are the D. speciosa of Dunker and of authors. Dunker gave the erroneous type locality as “Philippine Islands,” which is here corrected to Anaa Island, Tuamotu Islands. Reeve’s figure is based on a specimen in the Cuming collection from this locality. Nomenclature—Reeve in 1846 figured this spe- cies, calling it Ricinula clathrata Lamarck, vari- ety B, and both Crosse (1862) and Dunker (1867) in describing reeveana and speciosa respectively, cite Reeve’s figures in synonymy. However, R. reeveana, which is preoccupied (non C. B. Ad- ams, 1852), has been shown to be a synonym of D. rubusidaeus Réding, leaving Dunker’s name available for the present taxon. Records—TUAMOTU ISLANDS: (USNM; ANSP; FMNH; AMS); Garumaoa; Mataira; Oneroa; Opakea, all Raroia Atoll (USNM); Anaa Id. (MCZ); Vahitahi (USNM); Fangatau; Makatea (Dautzenberg and Bouge, 1933, p. 239). PITCAIRN ISLAND: Oeno Island (USNM). + Pearl f Hermes LL Is. iA 3 Christmes . es an © = ‘PHoEN 1s. . + TOKELAU IS. 4 + MARQUESAS IS. Same, “. v “830A ts NEW . gee + Moria Theresa tS (as rn ae Plate 26. Geographical distribution of Drupa (Ricinella) speciosa (Dunker). Open circles are literature records. INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 31 Drupa clathrata clathrata (Lamarck, 1816) (Pl. 2, figs. 16-18; Pls. 27, 28) Range—From Japan and the Philippines to the Marquesas and Pitcairn Islands. Remarks—The large number of spines, rugose columella and apertural coloration serve to char- acterize this species. Populations occurring in the Indian Ocean are separable on shell morphology and are'recognized as a subspecies, D. clathrata miticula (Lamarck). Habitat—On reefs, in crevices and under coral rocks, intertidal. Description—Shell 16 to 57 mm (% to 2% inches) in length, ovate and solid, spire short; body whorl large, crossed by five rows of spiral ribs. Ribs with siphonous, spiny tubercles; tubercles higher towards the margin of the outer lip. Spiral ribs connected by low axial ribs to form shallow pockets in the interstices. Surface sculptured with fine spirally arranged scales. Aperture sub- elliptical, columella excavated posteriorly, with 4 to 5 small plications above the siphonal canal. Parietal shield thinly enameled, reflected to form an irregular callus. Sutural canal well developed as a groove turned toward the spire. Axial fold strong, extending the length of the columella and terminating as a margin of the siphonal canal. Outer lip dentate with 4 to 5 singularly arranged teeth which sometimes become united. Color brownish-white externally, margin of aperture spotted with brown, interior of aperture a light violet. Operculum typical of group. The radula has the side-cusps of the rachidian bifid or trifid, and there are 2 to 3 deeply rooted lateral denticles, exclusive of the end-cusps. O.1MM =I Plate 27. Radula of Drupa (Ricinella) clathrata clathrata (Lamarck). Half a transverse row; Pango Point, Efate Id., New Hebrides. [14-845] 32. Ricinella Plate 28. Figs. 1, 2. Drupa (Ricinella) clathrata clathrata (Lamarck); Fig. 3. D. (R.) clathrata miticula (Lamarck). Fig. 1. Holotype of Ricinula clathrata Lamarck (MHNG no. 1101/14/1; 30.4 x 31.2 mm). Fig. 2. Specimen of Drupa (Ricinella) clathrata clathrata (Lamarck), from Pango Point, Efate Id., New Hebrides (WOC coll.; 28.3 x 25.8 mm). Fig. 3. Lectotype of Ricinula miticula Lamarck (MHNG no. 1101/13/1; 26.3 x 21.0 mm). Measurements (mm)—(including specimens with a mature lip) spines; all length width 57.3 53.0 “South Seas” 42.8 38.2 “Central Pacific” 30:1 29.0 Suva reef, Fiji Ids. 32.2 24.6 Samar Id., Philippines 30.4 31.2 Holotype of clathrata Lamarck 23.1 20.0 Tacume Id., Tuamotu Ids. 21.0 18.5 Pango Pt., New Hebrides Synonymy— 1816 Ricinula clathrata Lamarck, Tableau Encycl. Méthod- ique, p. 2, pl. 395, figs. 5a, b (no locality given); 1822 Lamarck, Hist. nat. anim. s. vertébres, vol. 7, p. 231; 1846 Reeve, Conchologia Iconica, vol. 3, pl. 2, fig. 9b only (Elizabeth Id., = Tuamotu Ids.); 1859 Chenu, Manuel de Conchyliologie, vol. 1, p. 168, textfig. 816; 1933 Dautzenberg & Bouge, Journal de Conchyliologie, vol. 77, p. 237. Emerson and Cernohorsky Muricidae 1835 Purpura clathrata Lamarck, Kiener (pars), Spéc. gén. icon. coq. viv., vol. 8, p. 15, pl. 3, fig. 5 only. 1853 Pentadactylus clathratus Lamarck, H. & A. Adams, Gen. Rec. Mollusca, vol. 1, p. 130; 1875 Troschel, Gebiss d. Schnecken, vol. 2, p. 133, pl. 13, fig. 3 (radula). 1880 Ricinula hystrix var. clathrata Lamarck, Tryon, Manual Conchology, vol. 2, p. 184, pl. 56, figs. 197, 198. 1913 Drupa rubuscaesia Bolten, Hedley (pars), Nautilus, vol. 27, no. 7, p. 80 (non D. rubuscaesius Réding, 1798). 1936 Drupa rubuscaesia Réding, Hirase, Coll. Jap. shells, p. 79, pl. 110, fig. 10; 1957 Kaicher, Indo-Pacific Sea Shells (Muricacea, Buccinacea), pl. 3, fig. 15 (non D. rubuscaesius Réding, 1798). 1951 Drupa rubuscaesius Réding, Hirase & Taki, Handb. illust. shells colour, pl. 110, fig. 10; 1959 Kira, Col. illust. shells of Japan, vol. 1, p. 58, pl. 23, fig. 9 (non Réding, 1798). 1954 Drupa rubuscaesium Réding, Kira, Col. Ilust. shells of Japan, pl. 23, fig. 9 (non D. rubuscaesius Réding 1798). 1962 Drupa rubsidaeus (sic) Réding, Kira, Shells west. Pa- cific in colour, p. 63, pl. 24, fig. 9 (non D. rubusid- aeus Réding, 1798). 1965 Drupa rubusidaeus Roding, Arakawa, Venus: Jap. Journ. Malacology, vol. 24, no. 2, p. 115, pl. 13, fig. 7 (radula) [non Réding, 1798]. 1967 Drupa_ (Ricinella) rubusidaeus (Roéding), Habe & Kosuge, Stand. book Jap. shells in color, vol. 3, p. 70, pl. 27, fig. 29 (non Roding, 1798). 1969 Drupa clathrata (Lamarck), Cernohorsky, Veliger, vol. 11, no. 4, p. 298, pl. 47, fig. 6. Types—The holotype of Ricinula clathrata Lamarck, is in the Muséum d’Histoire Naturelle, Geneva, no. 1101/14/1 (PI. 28, fig. 1). According to Rosalie de Lamarck’s marginal annotations in her father’s copy of the “Histoire naturelle des animaux sans vertebres,” only a single specimen was present in Lamarck’s collection at the time of description. In 1822 Lamarck gave the size of his specimen as 13% lignes [= 30.4mm], and this dimension agrees with the larger specimen, which is considered the holotype, but not the smaller 29.7mm specimen which accompanies it. This latter specimen has probably been added at a later date. The type locality here designated is Tuamotu Islands (after Reeve, 1846), specifically Raroia Island (from which there are specimens in the ANSP). Nomenclature—This species seems to have been confused with Drupa rubuscaesius Roding, and D. rubusidaeus Réding, by modern authors. Hedley (1913) initiated the confusion by suggest- ing that Ricinula clathrata Lamarck, R. speciosa Dunker and Purpura spathulifera Blainville, were synonyms of Drupa rubuscaesius Roding. Kira (1954, pl. 23, fig. 9; 1962, pl. 24, fig. 9) illus- trates a specimen of Drupa clathrata with an im- mature lip and calls it in the first instance D. rubuscaesium Roding, and in the second in- stance D. rubsidaeus (sic) Roding. Kaicher (1957, [14-846] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa_ 33 Plate 29. Geographical distribution of Drupa (Ricinella) clathrata clathrata (Lamarck) and its subspecies D. (R.) clathrata miticula (Lamarck). Open circles are literature records. pl. 3, fig. 15) cites D. clathrata as a synonym of D. rubuscaesia Roding, although it is not clear to which species her figures refers. As discussed earlier, Drupa rubuscaesius Roding, is a synonym of D. ricinus (Linnaeus), for Réding refers to Martini’s figure of the yellow apertured form of that species. Records—PHILIPPINES: Gigmoto, Catanduanes Id. (AMNH; DMNH; AIM); Borongan village, E. Samar Id. (ANSP); Marinduque Id. (USNM). FORMOSA (Taiwan): Chaiting (Janowsky coll.). RYUKYU ISLANDS: Yakushima: Onna Beach, N. of Naha, Okinawa; Bolo Point, Zampa Mis- aki, Okinawa (all USNM). JAPAN: Hackijo Id., 275 mi S. of Tokyo; Tosa, Shikoku; Kagoshima, Kyushu (all ANSP); Oshima, Osumi (USNM). BONIN ISLANDS: Chichi Shima (ANSP); Ani Jima (USNM). MARIANAS: Lagunan Tanapag, Saipan Id.; Ngargersul Id. (both ANSP); off Leprosarium, SW Tinian Id. (MCZ); Aspurguan, Guam Id.; Ypao Point, Maug Id. (USNM). WAKE ISLAND: (AMNH). MARSHALL ISLANDS: (many atolls, see map); Eniwetok: Bikini; Ronge- lap; Kwajalein; Jaluit (all USNM). SOLOMON ISLANDS: Choiseul Bay, Choiseul Id. (AMNH). NEW HEBRIDES: Pango Point, Efate Id., (AIM). LOYALTY ISLANDS: Lifu (USNM; AMS). NEW CALEDONIA: Touho (AMNH). GIL- BERT ISLANDS: Kingsmill Id. (USNM). FIJI ISLANDS: Cuvu Beach, S. Viti Levu (AMNH); Rat Tail Passage, Suva reef, S. Viti Levu (WOC coll.); SE of Onea Driki, Lau group (USNM). TONGA ISLANDS: Ha’ateiho reef, Tongatapu; Niuafo’ou (both USNM). NIUE ISLAND: (DMNH); Oneone reef; Utuko reef (both AMNH): Alofi (USNM). SAMOA IS- LANDS: Tutuila Id. (ANSP); Swan’s Id. (MCZ). Lalomalava, Savaii (DMNH). COOK ISLANDS: Bird’s Id., Palmerston Atoll; North Id., Palmerston Atoll (both USNM); Avaavaroa Passage, S. Rarotonga (ANSP). SOCIETY ISLANDS: Moorea; Venus Point, Tahiti (both AMNH); Punaavia, Tahiti (ANSP). TUAMOTU ISLANDS: Tacume: Raroia (both ANSP); Anaa Id. (AMS); Makemo: Tikahau; Vahitahi; Maka- tea (all USNM). MARQUESAS ISLANDS: Atuona Bay, Hiva- Oa Id. (ANSP). PITCAIRN ISLAND: (AMNH). LINE IS- LANDS: Caroline Id. (ANSP); Jarvis (DMNH). Drupa clathrata miticula (Lamarck, 1822) (Pl. 2, figs. 19, 20; Pls. 28, 30) Range—Madagascar to the Island of Timor in Indonesia. Remarks—First described by Lamarck in 1822, the identity of this Indian Ocean form appears to have escaped notice for over a hundred years and in that interval has been recorded as typical Drupa clathrata. It lacks the brown coloration of the columella and inner and outer lips that char- acterize the nominate subspecies. D. miticula is also a smaller shell, with shorter spines. The In- dian Ocean populations have a more pronounced purple color in the aperture than those of the nominate subspecies which is lavender to whitish purple. The exterior lacks the black spines of D. morum. Habitat—In tide pools, basalt rock, weed and some coral, and on sand and grass, from 0 to 8 feet. In the Cocos-Keeling Islands the species was found in “strong surf on a large boulder on the northern seaward reef at Horsburgh Island” (V. Orr Maes, personal communication). On Christmas Island it was found among Caulerpa I 0.1 mm Plate 30. Radula of Drupa (Ricinella) clathrata miticula (Lamarch). Half a transverse row; Greta Beach, Christmas Id., Indian Ocean. [14-847] 34. Ricinella Emerson and Cernohorsky mats and in rock and tide-pools (leg. A. Slack- Smith and A. Patterson). Description—Shell 16 to 38 mm (% to 12 inches) in length, ovate, globose, spire short, acuminate. Body whorl large with six transverse rows of short tubercles. Interstices between tubercles striated with four to six rows of closely spaced granules. Aperture moderately wide, oval, over three-quarter of shell length. Outer lip crenulated between tubercles, inner margin dentate with six small white teeth from which six conspicuous white raised lines run into the aperture. Inner lip enameled with three to four plait-like ridges pro- jecting into the aperture. Columella excavated posteriorly. Posterior siphonal canal open, obliquely curved toward the apex. Color pale brown on the exterior, interior of aperture purple. In the radula examined, one of the side-cusps of the rachidian was trifid while the other one was simple; there were 2 lateral denticles apart from the end-cusps. Measurements (mm)—(including spines; all specimens with a mature lip). length width 38.0 31.3 Christmas Id., Indian Ocean 34.3 28.0 S. Mahébourg, Mauritius 32.5 26.0 S. Mahébourg, Mauritius 26.3 21.0 Lectotype of miticula Lamarck 17.0 13.8 Christmas Id., Indian Ocean 16.5 14.5 Souillac, Mauritius Muricidae Synonymy— 1822 Ricinula miticula Lamarck, Hist. Nat. anim. s. verte- bres, vol. 7, p. 231 (no locality given); 1832 Blain- ville, Nouv. Ann. Mus. d’Hist. Nat. Paris, ser. 3, vol. 1, p. 211; 1844 Deshayes & Milne-Edwards, Hist. nat. anim. s. vertebres, ed. 2, vol. 10, p. 48 (refers to Kiener, fig. 5, junior = fig. 5a, 5a). 1835 Purpura clathrata Lamarck, Kiener (pars), Spéc. gén. icon. coq. viv., vol. 8, p. 15, pl. 3, figs. 5a, 5a (non Ricinula clathrata Lamarck, 1816). 1968 Drupa clathrata Lamarck, Taylor, Phil. Trans. Roy. Soc. London, ser. B, vol. 254, p. 201 (non Ricinula clath- rata Lamarck, 1816). Types—Two syntypes of Ricinula miticula are in the Muséum d Histoire Naturelle, Geneva. The slightly larger specimen, length 26.3 mm _ (PI. 28, fig. 3), no. 1101/13/1, is here selected as the lectotype of R. miticula. No type locality was given, and Mahébourg, Mauritius, is here desig- nated as the type locality (specimens in AMNH). Nomenclature—Kiener (1835) regarded _ Rici- nula miticula of Lamarck to be a juvenile of R. clathrata, but his figures are typical R. clath- rata miticula. Records—(Specimens): MAURITIUS: Gris Gris, 1 mi. ESE of Souillac; Vacoas Point, 3 mi. S. of Mahébourg; Point Pimente, N. side Arsenal Bay; Pointe Fayette; Caves Point (all ANSP); Mahébourg (AMNH); near Port Louis (MCZ). COCOS-KEELING ISLANDS: N. end of Horsburgh Id (ANSP). CHRISTMAS ISLAND: Lily Beach; Greta Beach; Dolly Beach (all WAM). INDONESIA: Timor (AMS). Records—(Literature—identified as “clathrata”): SEY- CHELLES ISLANDS: Coetivy Id. (Melvill, 1909, p. 104); Mahé (Taylor, 1968, p. 201). MADAGASCAR: (Dautzenberg, 1923, p. 38). MAURITIUS: (Viader, 1937, p. 32). REUNION ISLAND: (Deshayes, 1863, p. 115). [14-848] May 30, 1973 Subgenus Drupina Dall, 1923 Type: Drupa grossularia Réding, 1798 Shell sub-ovate, strong, heavy, flattened dorso- ventrally, spire short; whorls ribbed spirally with inconspicuous nodules, surface sculptured with minute imbricated scales; columella doubly _pli- cated axially, outer lip with two well developed marginal processes. Operculum typical for genus. Radula with a broad but low rachidian which has from 13-18 cusps; the central cusp may be large or small and the 2 flanking side-cusps are tridentate and usually smaller than the central cusp. The lateral teeth are small and_ slender, with a smaller and more rounded base than in Drupa sensu stricto. The 2 members of this subgenus have a por- tion of the outer lip expanded as two conspic- uous lobate processes. The process develops as the individual reaches maturity and thus is not laid down and re-absorbed by the mantle as the shell grows; instead the processes are thickened, extended and often bifurcated. Both species of the subgenus Drupina are Indo-Pacific in dis- tribution. Synonymy— 1923 Drupina Dall, Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 75, p. 303. Type-species by original designation: Ricinula digitata Lamarck, 1816 [= Drupa grossularia Ré6ding, 1798). | Ochi Plate 31. Radula of Drupa (Drupina) grossularia Roding. Half a transverse row; Fiji Islands. INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 35 Drupa grossularia Réding, 1798 (Pl. 2, figs. 23, 24; Pls. 31, 32) Range—From the Cocos-Keeling Islands in the Indian Ocean to West Australia and throughout the Pacific to Hawaii and the Marquesas Islands. Remarks—This is a very distinctive species with the large digitate processes and solid yellow aperture distinguishing it from all other mem- bers of the genus except Drupa lobata (Blain- ville), a dark brown apertured form inhabiting the Indian Ocean. Habitat— Lives clinging to rocks exposed at low tide or ina few feet of water on windward rather than leeward reefs. Demond (1957) records a specimen taken alive at a depth of 32-38 feet in a lagoon west of Saipan. Description—Shell 18 to 33 mm (% to 14 inches) in length, ovate, spire very short, body whorl large. Whorls spirally ribbed with low rounded nodules most common; nodules often as siphon- Plate 32. Drupa (Drupina) grossularia Réding. Fig. 1. Lectotype figure of Drupa grossularia Réding, from Martini, 1777, Syst. Conchylien-Cabinet, vol. 3, pl. 102, fig. 978. Fig. 2. Type figure of Purpura laurentiana Petit de la Saussaye, from Journal de Conchyliologie, 1850, vol. 1, pl. 13, fig. 2; Pacific Ocean (20.0 x 16.0 mm-—yjuvenile specimen). Fig. 3. Lectotype of Ricinula digitata Lamarck (MHNCG no. 1101/16/1; 22.4 x 22.0 mm) [14-859] 36 = Drupina ous tubercles near the margin of the aperture. Sculpture of crisp, well-defined scales; detail of sculpture retained only in well preserved indi- viduals. Aperture in adult stage, linear, con- stricted by 5 or 6 singularly arranged, close-set teeth projecting from the outer lip, and a col- umellar callus with from 2-5 inconspicuous pli- cations. Columella doubly folded axially. Aper- ture of juvenile specimens expanded. In adults, siphonous, digitate processes extend from the first and second ribs of the body whorl. The proc- esses tend to bifurcate at maturity; canal of the upper process canaliculate, lower canal generally sealed in the adult stage. Exterior of shell white or cream, aperture yellow to orange, denticles of outer lip white, columellar plications faintly whit- ish. Operculum typical for the genus, dark orange-brown in color. The radula is of a considerably modified dru- pine type and has been described in the sub- generic diagnosis. The size and length of the cen- tral tooth and accessory lateral denticles are quite variable. Measurements (mm)—(including digitations; all specimens with a mature lip). length width 32.7 33.6 Limu, Niue Island 32.0 30.9 S. Luzon Id., Philippines 23.2 22.2 Kavieng, New Ireland 22.4 22.0 Lectotype of digitata Lamarck 18.7 18.3 Bougainville, Solomon Ids. Synonymy— 1685 Lister, Hist. Syn. Meth. Conchyliorum, pl. 804, fig. 12. 1758 Seba, Locupl. rer. nat. thes. descriptio, vol. 3, pl. 60, fig. 48. mon « binomial). 1791 Murex neritoideus Gmelin (pars), Systema Naturae, ed. 13, p. 3537 (refers to Seba, op. cit., Lister, op. cit., and Martini, figs. 978, 979 only) [non Linnaeus, 1767]. 1798 Drupa grossularia Réding, Museum Boltenianum, p. 55 (refers to Martini, op. cit.) [no locality given]; 1913 Hedley, Nautilus, vol. 27, no. 7, p. 80; 1957 Kaicher, Indo Pacific Sea Shells (Muricacea, Buccinacea), pl. 4, fig. 4; 1970 Salvat, Cahiers du Pacifique, no. 14, p. 46. 1816 Ricinula digitata Lamarck, Tableau Encyclopédique Méthodique, p. 2, pl. 395, figs. 7a, b (no locality given); 1822 Lamarck, Hist. nat. anim. s. vertébres, vol. 7, p. 232; 1827 Crouch, Ilust. Introd. Lamarck’s Conchology, p. 36, pl. 18, fig. 8; 1842 Reeve (pars), Conchologia Systematica, vol. 2, p. 215, pl. 256, fig. 3 only; 1846 Reeve, Conchologia Iconica, vol. 3, pl. 1, fig. 2a (Lord Hood Id. = S. Marutea Id.); 1859 Chenu, Manuel de Conchyliologie, vol. 1, p. 168, text fig. 815; 1880 Tryon, Manual of Conchology, vol. 2, p. 185, pl. 56, fig. 191 and pl. 57, fig. 203; 1933 Dautzenberg & Bouge, Journal de Conchyliol- ogie, vol. 77, p. 237. Emerson and Cernohorsky Muricidae 1842 Purpura monstruosa Lesson, Rev. Zool Cuvierienne, vol. 5, App. p. 103 [Gambier Islands]. 1823 Murex fimbriatus Mawe. Linn. Syst. Conchology, p. 131, pl. 26, fig. 4 (non Brocchi, 1814; nec Lamarck, 1822). 1825 Murex ricinus Wood, Index Testaceologicus, pl. 26, fig. 5Sla (non Linnaeus, 1758). 1832 Purpura digitata Lamarck, Blainville, Nouv. Ann. Mus. (Hist. Nat. Paris, ser. 3, vol. 1, p. 210; 1833 Quoy & Gaimard, Voyage L’Astrolabe, vol. 2, p. 578, pl. 39, figs. 20-22 (shell, animal and operculum) [Carteret Harbour, New Ireland]; 1835 Kiener, Spéc. gén. icon. coq. viv., vol. 8, p. 16, pl. 3, figs. 6, 6a. 1850 Purpura laurentiana Petit de la Saussaye, Journal de Conchyliologie, vol. 1, no. 4, p. 403, pl. 13, fig. 2 (Pacific Ocean) [juvenile specimen]. 1853 Pentadactylus grossularius Bolten, H. & A. Adams, Gen. Rec. Mollusca, vol. 1, p. 129 and vol. 3, pl. 13, fig. 6c; 1875 Troschel, Gebiss d. Schnecken, vol. 2, p. 133, pl. 13, fig. 1 (radula). 1880 Ricinula hystrix var. laurentiana Petit, Tryon, Manual of Conchology, vol. 2, p. 184, pl. 56, fig. 192 (juvenile specimen). 1908 Pentadactylus (Pentadactylus) digitatus Lamarck, Horst & Schepman, Cat. Syst. Moll. Mus. Hist. Nat. Pays- Bas, vol. 13, p. 157. 1929 Drupina grossularia Rodding, Iredale, Mem. Queens- land Museum, vol. 9, pt. 3, p. 290; 1961 Rippingale & McMichael, Queens]. & Gt. Barrier reef Shells, p. 102, pl. 13, fig. 2; 1962 Kira, Shells west. Pacific in color, p. 62, pl. 24, fig. 3; 1965 Arakawa, Venus: Jap. Journ. Malacology, vol. 24, no. 2, p. 116, pl. 13, figs. 8-10 (radula); 1969 Cernohorsky, Veliger, vol. 11, no. 4, p. 303, pl. 48, fig. 11 (shell), text fig.7 (radula); 1971 Wilson & Gillett, Australian Shells, p. 92, pl. 61, figs. 5, 5a; 1971 Kay, Pacific Science, vol. 25, pp. 263, 275. 1952 Sistrum digitatum Lamarck, Morris, Field Guide to shells Pacific coast and Hawaii, p. 187, col. pl. 5, fig. 5; pl. 39, fig. 3. 1965 Drupina glossularia (sic) (R6ding), Wu, Bull. Inst. Zool. Acad. Sinica, vol. 4, p. 99, text figs. 20, 21 (radula); 1967 Habe & Kosuge, Stand. Book Jap. shells in color, vol. 3, p. 70, pl. 27, fig. 22 (invalid emenda- tion). 1968 Drupa (Drupina) grossularia Réding, Orr Maes, Pro- ceedings of the Academy of Natural Sciences of Phil- adelphia, vol. 119, no. 4, p. 130. Types—The holotype of Drupa grossularia is no longer traceable and the specimen figured by Martini on plate 102, figs. 978, 979 (Pl. 32, fig. 1) which was cited by Réding, is here designated as the lectotype of the species. Two syntypes of Ricinula digitata Lamarck, are in the Muséum dHistoire Naturelle, Geneva, and the 22.4mm long specimen, no. 1101/16/1, which most closely corresponds to Lamarck’s cited dimen- sions, is here selected as the lectotype (PI. 32, fig. 3). The holotype of Purpura laurentiana Petit de la Saussaye, is in the Muséum National d’His- toire Naturelle, Paris (Journ. de Conchyliologie coll.). No type locality was given by Réding for D. grossularia, and the earliest record of Car- teret Harbour, New Ireland, by Quoy & Gaimard, [14-860] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 37 160° | | i J) Swe Gr a SD eneeneeear See EMEA cs eee en ES Se oe cr ts a |e ere | _| 2 ana Ravens —, | + Aohnatorr i) | ae et | | “ y & A HILIERINE ons eo . be ‘ Chppertor SSS niet so @ GROSSULARIA By J) Ws gw . e \ TRA AE | F z ; ie magi . Cicada feat a! | gp cietttaw = | o* Scie LS ae Galapagos i eae Cee % % ©@,,; be ¢ s 4 4 MARGUESAS IS. = Cocos - KeRLint re oe e Rory cs 1 ARC 3 -@e | 2 te * | : ie OO @| = = a 5 eae : ‘3 | | ae er ees ke eee ee : Jj ida eis faces | | | | Y | | | ee f { ae | | | a ZFALAND mm esa | \ | < | | 2 a 60" BO" -_ roe 20 y40" 160" : 180 Teo" 140 120" Too? 3° Plate 33. Geographical distribution of the species Drupa (Drupina) grossularia Réding, in the Pacific and East In- dian Oceans (full circles) and D. (D.) lobata (Blainville) in the Indian Ocean (half-open circles). 1833, is here designated as the type locality, as well as for Purpura digitata Lamarck. Two syn- types of Purpura monstruosa Lesson, which are said to be “typical” examples of D. grossularia, are in the Muséum National d’Histoire Naturelle, Paris (teste G. Richard). Nomenclature—There can be no doubt as to Réding’s concept of Drupa grossularia, as he, Schumacher and Lamarck, all cite identical figures in Martini, 1777. Petit’s Purpura laurentiana is a young specimen in which the mature lip has not started to form and has the whorls covered with a foreign growth (PI. 32, fig. 2). A similar juve- nile is illustrated on PI. 2, fig. 24. Records—COCOS-KEELING ISLANDS: S. end of Home Id. (ANSP). INDONESIA: Marudu Bay, N. Borneo (ANSP) Malawali Channel, N. Borneo (AMNH; USNM); Bali; Mor- otai Id. (both AMS); Oong Bay, Mandidi Id., Moluccas (MCZ); Pulau Pombo, Wasi, Ambon Id. (FMNH). PHIL- IPPINES: Recorded from many localities throughout the Archipelago: Luzon; Mindoro; Catanduanes; Busanga; Bala- bac; Sulu Archipelago (AMNH; ANSP; MCZ; USNM); Boron- gan village, E. Samar Id. (AIM). FORMOSA (Taiwan): Now Wow (USNM). RYUKYU ISLANDS: Sobe; Kadena; Mekan- iko; Onna reefs; Bolo Point, all Okinawa (all AMNH); Miyako (FMNH). JAPAN: Hachijo Id., 275 mi. S. of Tokyo; Tosa, Shikoku; Osima, Osumi (all ANSP). MARIANAS: Lagunan Tanapag, Saipan; Agat Bay, Guam (both ANSP); Tinian Id. (MCZ). PALAU ISLANDS: Helen reef, Koror; Babelthuap (both ANSP). CAROLINE ISLANDS: S. of Gar- abayo; Kayangel; Rattakadokoru; Ngaruangl; Ngangersul; Ponape; Yap (all ANSP); Elato Atoll; Lamotrek Atoll; Sata- wal Atoll; Kapingamarangi (all USNM); Lukunor (AMNH). MARSHALL ISLANDS: Eniwetok Atoll; Bikini Atoll; Kwaja- lein Atoll (all USNM); Arno Atoll (AMNH). ADMIRALTY ISLANDS: Manus Id. (DM). NEW BRITAIN: Rabaul (AMNH,; USNM). NEW IRELAND: Kavieng (AMNH). NEW GUINEA: Aoeri Ids., Geelvink Bay; SW Biak, Schouten Ids.; Wooi Bay, Japen Id.; 1 mi. NE Mioes Woendi, Padaido Ids. (all ANSP); Samarai, Papua; Port Moresby, Papua (both AMNH); Milne Bay, Papua (USNM). AUSTRALIA: Torres Strait (Shirley, 1912, p. 102); Queensland: Michaelmas Cay, off Cairns; Bramble Bay, off Lucinda (both AMS); West Australia: Barrow Id. (Wilson & Gillet, 1971, p. 92. SOLO- MON ISLANDS: Vanikoro Id., Santa Cruz group; Reef Id., Santa Cruz group (both AMS); Kieta, Bougainville Id.; Lutee, Choiseul Id.; Ataa, N. Malaita Id. (all AMNH); Ugi Id., Shortland group (USNM); Ticopia Id., (AIM); NEW HEB- RIDES: Meli Id., SW Efate Id.; Pango Point, Efate Id. (both AIM); Pentecost Id. (Powell coll.). LOYALTY ISLANDS: Lifu (AMS). NEW CALEDONIA: Touho (AMNH); Bourail; 7 mi. SW Gatope Id., Voh (both ANSP). FIJI ISLANDS: 3 mi. NE Tunuloa, Vanua Levu (MCZ); Suva Harbour, S. Viti Levu; Ogea Levu, Ogea, Lau group (both USNM); Wadigi Id., Mamanuca group (WOC coll.). GILBERT ISLANDS: Abaiang (MCZ); Onotoa Atoll, Kingsmill Ids. (USNM). EL- LICE ISLANDS: Funafuti (AMS; AIM). WALLIS & FUTUNA: Nukuhifala, Wallis Id.; W. coast of Uvea, Wallis Ids.; E. side of Faioa, Wallis Ids. (all USNM); Anse de Sigave, Hoorn Id., Futuna Ids. (USNM). SAMOA ISLANDS: Asau Harbour, Savaii (USNM):; Vailele Bay, Upolu Id. (ANSP); Satalo Id., Upolu Id. (AIM); Apia, Upolu Id. (Powell coll.); Pago Pago, Tutuila Id. (AMNH; MCZ). TONGA ISLANDS: Ha’ateiho, Tongatapu (USNM). NIUE ISLAND: Limu (AMNH); Alofi (USNM). PHOENIX ISLANDS: Enderbury Id. (USNM). COOK ISLANDS: Akamaru, Manihiki Atoll (ANSP); Bird’s Id. and Tom’s Id., Palmerston Atoll; Motu Akaiami, Aitutaki (all USNM); Mauke (ANSP); several localities on Rarotonga (MCZ; USNM; AIM). AUSTRAL ISLANDS: Rurutu; Raivavae (both USNM). SOCIETY ISLANDS: N. of Fare, Huahine; Faredine, NW Moorea; Papeete, Tahiti (all USNM). TUA- MOTU ISLANDS: Raroia Id.; Lord Hood Id. [= S$. Marutea Id.] (both AMNH); Makemo Id.; Toau Atoll (both ANSP); Nengonengo Id. (USNM). GAMBIER ISLANDS: Mangareva Id. (USNM). MARQUESA ISLANDS: (USNM; ANSP). LINE ISLANDS: Palmyra Id.; Flint Id.; Christmas Id. (all ANSP); Fanning Id.; Kingman reef; Washington Id.; Jarvis Id. (all Kay, 1971, p. 275). HAWAIIAN ISLANDS: Kure Id.; Midway Id. (both USNM; Makua, Oahu (Adams, 1967, p. 4). Cm nai L O.1 am J Plate 34. Radula of Drupa (Drupina) lobata (Blainville). Half a transverse row; Veeckens Bay, S. Pagi Id., Indonesia. [14-861] 38 = Drupina Drupa lobata (Blainville, 1832) (PI. 2, figs. 25, 26; Pls. 34, 35) Range—From the Red Sea and East Africa through the Indian Ocean to Thailand, Sumatra and West Australia. Remarks—Despite its closeness to Drupa gross- ularia, the present chocolate-mouthed species seems to be a distinct form which replaces the yellow-mouthed species in the Red Sea and In- dian Ocean. Their ranges are known to overlap in the Cocos-Keeling Islands where the two forms were collected at Home Island and no intermediates were found (Ostheimer and Orr Maes leg.), and along the coast of West Australia (specimens in West Australian Museum). Habitat—On intertidal wave-swept reefs. In the Cocos-Keeling Islands it was found on the tops and sides of rocks near shore on the seaward reefs, and on the seaward ends of passes (Orr Maes, personal communication). Description—Shell 18 to 32 mm (% to 1% inches) in length, very similar to Drupa grossularia Rod- ing, but differing in having a chocolate-brown aperture, a brownish dorsum, and in having a wider, more lobate digitate process extending from the first rib on the body whorl. This proc- ess shows no tendency to bifurcate as it does in D. grossularia, and the canal remains open in the adult. The radula is similar to that of D. grossularia; the side-cusps of the rachidian are short, broad and trifid, and are followed by 6 moderately deeply rooted lateral denticles. Measurements (mm)—(including digitations; all specimens with a mature lip) length width 33.2 33.0 Mauritius 28.6 27.8 Kiwengwa, Zanzibar 26.5 26.3 Syntype of dactyloides Schumacher 26.2 27.5 Eilat, Gulf of Aqaba 18.5 17.5 Zanzibar Synonymy— 1817 Ricinella dactyloides Schumacher, Essai nouv. systéme, p. 241 (refers to Martini, 1777, vol. 3, pl. 102, figs. 978, 979 = Drupa grossularia Riding) [no locality given] (nomen oblitum). 1823 Ricinula digitata Lamarck, Sowerby, Gen. Rec. foss. shells, pt. 18, pl. 235, figs. 3, 4 (no locality given) [non Lamarck, 1816]. Emerson and Cernohorsky Muricidae 1832 Purpura lobata Blainville, Nouv. Ann. Mus. d’Hist. Nat. Paris, ser. 3, vol. 1, p. 210, pl. 9, fig. 7 (no locality given); 1835 Kiener, Spéc. gén. icon. coq. viv., vol. 8, p. 18, pl. 3, fig. 7. 1842 Ricinula digitata var. Lamarck, Reeve (pars), Conchol- ogia Systematica, vol. 2, p. 215, pl. 256, fig. 4 only (non Lamarck, 1816). 1844 Ricinula digitata var. fusca “Sowerby,” Deshayes & Milne-Edwards, Hist. Nat. anim. s. vertebres, ed. 2, vol. 10, p. 53 (no locality given) [published in syn- onymy of R. lobata Blainville—refers to Sowerby, 1823, pl. 235, fig. 4] (non R. fusca Kiister, 1862). 1846 Ricinula digitata var. B. Reeve, Conchologia Iconica, vol. 3, pl. 1, fig. 2b (Seychelles Ids.) [non Lamarck, 1816]. 1880 Ricinula digitata var. lobata Blainville, Tryon, Manual Conchology, vol. 2, p. 185, pl. 57, fig. 205. 1896 Ricinula lobatus Blainville, Shopland, Journ. Bombay Soc. Nat. Hist., vol. 10, p. 220. 1903 Sistrum digitatum (var. lobata) E. A. Smith in Gardi- ner, Fauna & Geog. Maldive & Laccadive Archi- pelago, p. 609. 1919 Drupa digitata var. lobata Blainville, Cooke, Proc. Malac. Soc. London, vol. 13, pt. 4, p. 101 (descrip- tion of radula); 1937 Viader, Mauritius Inst. Bull. vol. 1, pt. 2, p. 32. 1913 Ricinula lobata Blainville, Hedley, Nautilus, vol. 27. no. 7, pp. 79, 80; 1922 Hornell, Madras Fish. Dept. Bull., no. 6, p. 217. 1950 Drupa (Drupina) grossularia lobata Blainville, Abbott, Bull. Raffles Museum, vol. 22, p. 80. 1956 Drupa (Drupina) lobata (Blainville), Franc, Ann. L’'Inst. Océanog. Monaco, N.S. 32, p. 37 (Ile Abulat, Red Sea); 1967 Orr Maes, Proc. Acad. Nat. Sci. Philadel- phia, vol. 119, no. 4, p. 130, pl. 11, fig. E. 1961 Drupa lobata Spry, Tanganyika Soc. Notes & Record, no. 56, p. 21, pl. 7, fig. 142. 1969 Drupina lobata (Blainville), Cernohorsky, Veliger, vol. 11, no. 4, p. 303 1970 Heinicke, Hawaiian Shell News, vol. 18, no. 7; p. 6, text fig.; 1971 Wilson & Gillett, Australian Shells, p. 92, pl. 61, fig. 4 (Pt. Cloates, West Australia). Types—The type specimen of Purpura lobata Blainville, is presumably in the Muséum National dHistoire Naturelle, Paris. Five probable syn- types of Ricinella dactyloides Schumacher, are in the Zoological Museum, Copenhagen. Four of these specimens have the letters “Sp” marked either in the aperture or on the dorsum, and these originated from the Spengler collection. One specimen is marked “Sch” [= Schumacher] in- side the aperture (PI. 35, fig. 1). The type: local- ity of D. lobata is here designated as Mogadiscio, Somalia. Nomenclature—When Schumacher described Ricinella dactyloides, his diagnosis consisted of only the three words “labio externo digitato”; for an illustration he referred to Martini’s figures 978, 979, which represent Drupa_ grossularia Réding. However, the extant and probable syn- types of Ricinella dactyloides are referable to the species Drupa lobata (Blainville), and Schu- macher’s taxon would in effect have 15 years [14-862] May 30, 1973 Plate 35. Drupa (Drupina) lobata (Blainville). Fig. 1. Probable syntype of Ricinella dactyloides Schu- macher; marked;‘Sch[umacher]” inside aperture (ZMC; 26.5 x 26.3 mm). Fig. 2. Probable syntype of R. dactyloides Schumacher; marked#‘Sp[engler]” inside aperture (ZMC; 26.8 x 25.8 mim). priority over Blainville’s. Since Schumacher’s name has not once been applied to a taxon as the valid name during the last 50 years, it is con- sidered to be an unused senior synonym. The taxon Purpura lobata Blainville, however, has been in general current use during the preceding INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 39 fifty years, and has been used by 5 different authors in 10 publications (see Declaration 43 of the ICZN; Bull. Zool. Nomencl., vol. 27, pts. 3/4, p. 135). Hedley (1913) suggested that the name fusca Deshayes & Milne-Edwards, 1844, be applied to the present species on thé mistaken belief that Blainville had proposed Purpura lobata for the yellow-apertured Drupa digitata (Lamarck) [= D. grossularia Roding]. Blainville (1832, p. 210) clearly states: “couleur d'un’ brun-marron en dehors et & la circonférence de louverture, blanche en dedans”; the type-figure given by Blainville (1832, pl. 9, fig. 7) is also an excellent representation of the dark brown apertured form, despite the lack of cited locality. Records—RED SEA: Eilat, Gulf of Aqaba, Israel(A. Hadar; K. Haim; AMNH; DMNH); He Abulat (Franc. 1956, p. 37); Jidda, Saudi Arabia (DMNH) GULF OF ADEN: Aden (Shop- land, 1896, p. 220). EAST AFRICA: 9 mi. N. of Mogadiscio, Somalia (ANSP); at 19 km marker, N. of Mogadiscio, So- malia (AMNH); Diani Beach, Kenya (Heinicke, 1970, p. 7); 15 mi. SSE of Dar-es-Salaam, Tanzania; 4 mi. ESE of Dar- es-Salaam, Tanzania (both MCZ); Mozambique City, Mozam- bique (ANSP). ZANZIBAR: Pange Id. Kiwengwa; Mangapivani (all ANSP). SEYCHELLES: Beau Vallon Beach, Mahé (ANSP). MADAGASCAR: Grande Recife, W. end of ship pier, Tuléar; Grande Recife, W. of airport, Tulear (both MCZ). REUNION: (Deshayes, 1863, p. 115). MAURITIUS: NW side of Tamarin Bay (ANSP). MALDIVE ISLANDS: Imma Id., N. Male Atoll; Fodiffolu Atoll; Ari Atoll (all ANSP). LACCADIVE ISLANDS: (Hornell, 1922, p. 217). THAILAND: Goh Phi Phi; Goh Huyong, Similan Ids. (both USNM). COCOS-KEELING ISLANDS: N. tip West Id.; S. end of Direction Id.; S$. end Home Id. (all ANSP). CHRIST- MAS ISLAND: (Tomlin, 1935, p. 79). INDONESIA: Pulau Siburu, N. of Sipora, S.W. Sumatra;Pulau Bai, Batu group, off Sumatra; W. shore Veeckens Bay, S. Pagi Id. (all USNM). WEST AUSTRALIA: W. of Ningaloo homestead, Pt. Cloates, 22°42’S and 113°39’E (WAM). [14-863] 40 Index Emerson and Cernohorsky INDEX TO DRUPA NAMES IN VOL. 3 NO. 13 The number following the name refers to the pagination found at the top of the page. The col- umn at right is the looseleaf pagination. Al new names are in bold face type. aesculus Réding, 9 albolabris Blainville, 20 album Montfort, 20 arachnoides Lamarck, 20 Azumamorula, 8 baylei Coss. & Lambert, 5 bollonsi Suter, 7 botroides Réding, 9 Canrena Link, 14 chamaemorus Réding, 9 Condonia Hertlein, 8 cornus Roding, 9 clathrata Lamarck, 31 dactyloides Schumacher, 38 digitata Lamarck, 36 Drupa Réding, 14 Drupina Dall, 35 Drupinae, 13 elegans Broderip & Sowerby, 24 fimbriatus Mawe, 36 fragun, Roding, 28 fusca Deshayes & Edwards, 38 glans Réding, 9 globosa Morch, 16 globosum Martini, 16 grossularia Réding, 35 hadari Emerson & Cernohorsky, 23 hannai Howe, 8 hericinus Martini, 28 hippocastanum Wood, 28 horrida Lamarck, 16 hystrix Linnaeus, 20 hystrix auctt., 28 iodostoma Lesson, 18 jodostoma Boettger, 19 14-809 14-824 14-824 14-824 14-808 14-805 14-807 14-809 14-818 14-809 14-808 14-809 14-845 14-862 14-860 14-818 14-859 14-817 14-828 14-860 14-842 14-862 14-809 14-820 14-820 14-859 14-827 14-808 14-842 14-842 14-820 14-824 14-842 14-822 14-823 laurentiana Petit de la Saussaye, 36 lobata Blainville, 38 mancinella Roding, 9 miticula Lamarck, 33 monstrosa Lesson, 37 morum Roding, 15 muricina Réding, 9 mutica Lamarck, 8 neritoidea Link, 16 neritoideus Linnaeus, 16 neritoideus Mawe, 20 nodosa Linnaeus, 16 nodus Gmelin, 28 Pentadactylus Morch, 14 pisolina Lamarck, 8 purpurata Schumacher, 28 Purpurinae, 13 puruensis K. Martin, 5 reeveana Crosse, 28 rhombiformis K. Martin, 5 Ricinella Schumacher, 27 Ricinula Lamarck, 14 Ricinulus Demarest, 14 ricinus Linnaeus, 19 rubuscaesius Réding, 20 rubuscestus Dall, 20 rubusidaeus Roding, 27 Sistrum Montfort, 14 smithi Brazier, 8 spathulifera Blainville, 28 speciosa Dunker, 30 styriaca Stur, 5 Thaidinae, 13 trapa Réding, 9 tribulus Roding, 20 uva Réding, 9 violacea Schumacher, 16 vitiensis Pilsbry in P. & B., 9 walkerae Pilsbry & Bryan, 9 [14-864] Muricidae 14-860 14-862 14-809 14-847 14-861 14-819 14-809 14-808 14-820 14-820 14-824 14-820 14-842 14-818 14-808 14-842 14-817 14-805 14-842 14-805 14-841 14-818 14-818 14-823 14-824 14-824 14-841 14-818 14-808 14-842 14-844 14-805 14-817 14-809 14-824 14-809 14-820 14-809 14-809 INDO-PACIFIC WILLIAM H. DALLE MOLLUSCA —oSsiii'SEEs volume 3, no. 14 THIS SECTION CONTAINS 1. The Genus Gabrielona (Phasianellidae) in the Indo-Pacific and West Indies. By Robert Robertson. (Put in Binder 1, just after the guide tab “Pha- sianellidae” bo . Title page to Volume 3. (put in Binder 1). 3. List of Issues published as of June 1, 1973. (put in Binder 1). 4. Replacement Pages for Tectarius (8 pages of corrections for “The Family Littorinidae in the Indo-Pacific. Part I.” Put these 4 sheets in their proper place in Binder | by following the looseleaf page numbers at the bottom of the page (i.e. [05-443]). Remove and place the obsolete pages at the end of Binder 3, just after the guide tab “Replaced Pages.” This instruction sheet, issued June 1, 1973, may be destroyed, or placed at the end of Binder 3. May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 Gabrielona 4) THE GENUS GABRIELONA (PHASIANELLIDAE) IN THE INDO-PACIFIC AND WEST INDIES by RosBert ROBERTSON Pilsbry Chair of Malacology The Academy of Natural Sciences of Philadelphia This is the first part to be published of a mono- graph on the systematics of all the Indo-Pacific Phasianellidae. The other parts will comprise a general introduction, methods, a_ bibliography, index, and treatments of the genera Phasianella and Tricolia (including the subgenus Hiloa Pils- bry). Conventional approaches to the systematics of these groups, i.e. studies of shells and radulae, have yielded inconclusive results on how species should be categorized, and further biological studies are planned on them. The data on Gab- rielona systematics are published in the mean- time because they involve fewer unresolved prob- lems and because there are new taxa (G. pisinna, G. raunana goubini and G. sulcifera). Biological data relevant to systematics are unlikely to be obtained for this rarely collected genus. Small, low-spired Tricolia specimens, which are far more common than Gabrielona, have been mistaken for Gabrielona. The generic characters detailed in this paper should help to dispel such misidentifications in the future. Gabrielona is distantly related to the other two genera grouped in the Phasianellidae. Genus Gabrielona Iredale, 1917 Type-species: Phasianella nepeanensis Gatliff and Gabriel, 1908 Chief distinguishing characters—The shells are small (G. hadra, the largest known species, can be 3.3 mm. long), with globose outlines and low Plate 36. Gabrielona nepeanensis (Gatliff and Gabriel). Figs. 1:2, 5, 7-9. South Australia. Figs. 3-4, 6, 10. Victoria, Austra- lia. Fig. 10. Holotype. Fig. 1, x30; Figs. 2-10, x20. [04-151] 42. Gabrielona spires. Sculpture, if present, is most prominent on the early whorls. Colors are pinks, reds, or browns, and white. Inside the aperture there is a spiral palatal sulcus which, if high in the aper- ture, is positioned beneath a spiral, subsutural feature of the color pattern on the external sur- face. All the species but G. raunana have from one to three apertural denticles. All the species are umbilicate, with an umbilical channel. From shells of other Phasianellidae of generally similar shape and size, Gabrielona is most read- ily recognized by its operculum which, external- ly, has a pronounced marginal ridge and a con- cave central region. The other two genera have opercula that, externally, are convex. When fully withdrawn into the aperture, the operculum of Gabrielona fits shallowly against an axial ridge (or series of faint ridges); the opercula of the other genera fit more deeply against similar, but generally faint, ridges. The axial ridge of Gab- rielona very rarely is as deep in the aperture as shown in Pl. 52, fig. 3, ar. Gabrielona radulae are highly distinctive. Taxonomic history—When Iredale proposed Gabrielona, he did “not think it has really any close relationship with Phasianella,” even though suggesting that it “may be classed for the pres- ent in the family Phasianellidae,” and “judging from the shell characters and the operculum and dead animal of the [unnamed] Lord Howe spe- cies ... a Naticoid affinity suggests itself.” Characteristically, Iredale did not describe the genus or mention any distinguishing characters other than the naticoid operculum. Retaining Gabrielona in the subfamily Phas- ianellinae, Thiele (1929) stated it to be doubt- fully distinct from Chromotis H. and A. Adams, 1863. Following their description of what later came to be the type-species of Gabrielona, Gat- liff and Gabriel (1908) had mentioned a similar- ity to Phasianella neritina Dunker—the type-spe- cies of Chromotis. Wenz (1938) retained Gabriel- ona as a tentative synonym of Chromotis, which he ranked as a subgenus of Tricolia. Australian malacologists have disregarded Thiele and Wenz, and have followed Iredale in treating Gabrielona as a distinct genus, but without giving reasons. Finding that Gabrielona definitely is not a syn- onym of Chromotis (here considered a synonym of Tricolia), I re-established it, accorded it gener- ic rank, and described it for the first time (Rob- ertson, 1958). My study of the radula of “G. brevis (Orbigny)” [= G. sulcifera Robertson]— not the type species—revealed that it is rhipido- glossate but different from that in other Phasi- Robert Robertson Phasianellidae anellidae. Thus, Gabrielona does not belong in the Naticidae or any other group with taenio- glossate radulae. Absence of a nacreous internal shell layer, and possession of a calcareous oper- culum necessitated my retaining Gabrielona in Phasianellidae, where I arbitrarily placed it in the subfamily Tricoliinae. Relationships— The relationships of Gabrielona to other supraspecific taxa in the Phasianellidae are obscure. The resemblances with most other phasianellids—shells of small size with bright col- ors and complex and variable patterns—seem superficial. In view of the distinct apertural, op- ercular and radular characters, the genus may not even belong in the family. However, pending more thorough knowledge of relationships within the Trochacea, it seems best to continue to retain Gabrielona in the Phasianellidae. Shell description (abbreviations refer to the drawings)—Attains lengths of 1.1-3.3 mm., widths of 1.1-3.1 mm., and 3.0-4.6 whorls; spire angles 90°-125°; outlines globose; fairly thin to thick, and transparent to opaque. Protoconchs (p) in- sert to slightly exsert, slightly inflated, smooth or with a spiral keel, white, either not demarcated from teleoconch or slightly to fairly prominently demarcated, 0.9-1.2 whorls when demarcated. First whorls 0.20-0.29 mm. in diameter. Subse- quent whorls: inflated, rounded in profile; su- ture slightly impressed. Sculpture predominantly smooth, or with axial plicae or spiral keels, cords and sulci; if present, sculpture most pronounced on second whorl, gradually declining and lacking near periphery or wholly lacking on last whorl of large shells. Periostracum not observed, but very thin layer possibly present. Colors: pinks, reds, or browns, and white. Patterns various, but com- monly with subsutural, subperipheral, and _peri- umbilical discordances. Aperture roundly pyri- form; outer lip never thickened or everted, thin to thick; callus on upper parietal area thin to thick; a faint to fairly prominent axial ridge (ar) or series of faint ridges within the aperture (against which the fully withdrawn operculum abuts); the ridges fairly variable in position, but never distant from the edge of the outer lip; a fairly faint to fairly pronounced spiral palatal sul- cus (ps) high in aperture or near middle, com- monly positioned beneath a color pattern feature on external surface and terminating at apertural ridge. Depending on the species, 0-3 faint to rather prominent apertural denticles present, namely 1 palatal and opposite the shoulder in position (pd), 1 on the lower part of the colu- mella (cd,)—both these on the apertural ridge— [04-152] May 30, 1973 and 1 on the middle part of the columellar lip (cd,); in species with a lower columellar denticle (cd,), a faint, broad ridge spirals up the columel- la; apertural ridge, palatal sulcus, and apertural denticles fairly faint to lacking on small shells. Channel extending into umbilicus (uc) bordered on right by the outer edge of the columellar lip (a slope or an escarpment), and on the left (at an acute angle) by a fairly faint to prominent es- carpment arising from the outer edge of the lower half of the columellar lip (abnormally, es- carpment absent); umbilicus narrow to wide. Opercula— Unlike those of the two other genera in the Phasianellidae, the opercula of Gabrielona are externally concave. In view of the extreme rarity of live-collected Gabrielona in collections, it is convenient that these distinctive opercula are retained—albeit infrequently—in the apertures of some empty shells. The fresh opercula are fairly transparent and white. They all are pauci- spiral and have fine, irregular spiral and radial growth lines and wrinkles on the external and in- ternal surfaces; this fine sculpture is not shown in the accompanying drawings except where spe- cially prominent. The external surface has a prominent spiral ridge near the edge (except the non-spiral colu- mellar sector) and a flattish but slightly concave central region which can have a differently tex- tured or sculptured outer, spiral area. On all Gabrielona opercula, a callus of varied thickness overlays the upper two-thirds or three-quarters of the central region. This callus covers all or most of the spiral suture, is thickest near the middle columellar edge of the operculum, and its lower margin (marked c¢ on the drawings) is irregular and variable in position and height, with or with- out a distinct escarpment. Small opercula and the single one known of G. sulcifera have a thin or very thin callus and therefore the suture is clear- ly visible externally—as well as in transparency. In the different species, the spiral ridge and cen- tral region are variously sculptured. The non- spiral columellar edge of the operculum is bev- eled. The internal surface of the operculum is rela- tively flat, but the central part of the last whorl near the columellar margin can be slightly con- cave. The early whorls are slightly raised, and a low, spiral escarpment is at the suture. The cen- tral area (at the axis) on all the species with op- ercula available (G. nepeanensis, G. pisinna, G. raunana [both subspecies] and G. sulcifera) is circular, crested at the perimeter, and 0.11-0.14 mm. in diameter—i.e., correlates neither with the INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 Gabrielona 43 various diameters of the first whorl of the shell, nor with the maximum shell size attained by the various species. This circular central area can be conspicuously smoother, shinier, and more trans- parent than the remainder of the internal sur- face, and always bears a low, central boss. The operculum of G. raunana (both subspecies) dif- fers from those of the other species in having structural radial lamellae near the edge. Muscle attachment scars are never clearly defined. When fully withdrawn into the aperture, the operculum abuts against the apertural ridge (ar), and fits closely against the columellar lip as well as the inner surface of the outer lip. In those species possessing the two columellar den- ticles, the operculum pivots against the columel- lar lip between these. The slight differences be- tween species in the form of the columellar lip account for the slight differences in the outline of the columellar margin of the opercula. Anatomy—The very few live-collected speci- mens available all were preserved dry, so that the bodies were dried out and appressed against the outer shell wall in the aperture. In G. pis- inna, the gut—distended with whitish, calcareous fragments—was conspicuous, fairly short and U- shaped, arising at the left, extending posteriad and bending to the right at a position about half of a whorl back from the outer lip, and then ex- tending forward to the anus on the right near the outer lip. There was no indication of an enlarged stomach. The mantle edge appeared to be rather thick, but the total volume of the dried body seemed remarkably small. Jaws were not detect- ed but could be present. Radulae (Pls. 42 and 57)—I have been able to study the radulae of only G. pisinna and G. sul- cifera. These differ greatly from other known phasianellid radulae and also from each other. The specimens from which the radulae of the two species were extracted were collected about 9,200 miles apart (New Caledonia and Antigua), and the size difference between their shells is considerable: the shell volume of G. pisinna is about 19-fold smaller than that of G. sulcifera. Relative to shell size, the radula of G. pisinna is rather longer and considerably wider than that of G. sulcifera. These differences in actual and rela- tive sizes perhaps account for some of the great differences. In particular, the small absolute size (but not the greater relative width) of the radula in G. pisinna could account for the fewer margin- als and laterals (3 laterals instead of the 5 in G. sulcifera), and the more curved transverse rows of teeth. The “central” of G. pisinna possibly is a [04-153] 44 Gabrielona pseudocentral comprised of one or two pairs of the original innermost laterals fused together (perhaps also with the original central). The size differences could not account directly for some of the other interspecific differences: the multicusped laterals of G. pisinna and, in G. sulcifera, the unwinged central and laterals, the differently-shaped bases of the laterals, and the massive innermost marginals. The radular differences help to confirm the conclusion reached independently from study of the shells and opercula of the two species, name- ly that they are distantly related congeners. Lack- ing information as to the total diversity of Gab- rielona radulae, their possible ontogenetic changes, the structural consequences of their ab- solute and relative sizes, and bearing in mind the evolutionary plasticity of Tricolia radulae, I con- sider the radular differences inadequate evidence for separating G. sulcifera from G. pisinna in a different subgenus or genus. Relationships within genus—Related pairs of Recent taxa are all allopatric. Only in New Cal- edonia is more than one species of Gabrielona known to occur, and these (G. pisinna and G. raunana) are distantly related congeners. The most closely related taxa distinguished here are G. raunana raunana and G. raunana goubini, which are ranked as subspecies. G. pis- inna is a dwarf, tropical Indo-Pacific homologue of southeastern Australian G. nepeanensis. G. sulcifera of the Caribbean perhaps is related, al- beit fairly distantly, to the western Pacific G. raunana. G. hadra is a clearcut fossil precursor of G. sulcifera. G. nepeanensis and G. pisinna seem distantly related to their congeners. Color and pattern variations—All the species are variable in coloration and pattern: G. rau- nana and G. nepeanensis especially so, and G. pisinna least of all. The range of coloration of Gabrieclona is narrower than in each of the other two phasianellid genera. Most of the color and pattern variation in Gabrielona is gradational, but a discontinuous color variation is treated under G. raunana goubini. Sexual dimorphism—Not detected conchological- ly. Fossil history—The only fossil species known certainly to belong in the genus is “Tricolia” hadra Woodring from the Bowden Formation (Middle Miocene or possibly Pliocene-Pleisto- cene) in Jamaica, West Indies. Distribution of Recent species—Until I transfer- red a West Indian species to the genus (Robert- son, 1958), G. nepeanensis (Gatliff and Gabriel), Robert Robertson Phasianellidae from southeastern Australia, was the only named species in the genus. An unnamed and cursorily studied Gabrielona, mentioned by Iredale (1917), was live-collected in the “sub-littoral” at Lord Howe Island (about 400 miles east of the coast of northern New South Wales, Australia). This locality, shown with a circle on Pl. 39, is be- tween the known distributions of G. nepeanensis and the two tropical Indo-Pacific species. Ire- dale’s specimens from Lord Howe were not lo- cated at the Australian Museum by Dr. D. F. McMichael in 1962 (letter to Dr. R. T. Abbott dated July 10). One new tropical Indo-Pacific species, G. pis- inna, and one new subspecies, G. raunana gou- bini, are described and named here, and the West Indian species to which I misapplied the name Phasianella brevis Orbigny is named G. sulcifera. Gabrielona quite possibly occurs in other trop- ical and subtropical faunal areas, such as West Africa and the Panamic Province. The four known Recent species occur almost exclusively along the coasts of continents and high islands; the only known exception is G. raunana raunana Ladd, a subspecies known only as subfossil shells from an atoll. Abundance—Very few live-collected specimens of any of the species have been available for study: only 7 G. pisinna and 1 G. sulcifera. The other two Recent species, including the type- species G. nepeanensis, are known only from empty, beach worn or subfossil shells, a few with opercula. Thus, the genus appears to be a relict group. Alternatively, it may have been rarely col- lected if, as I suspect, it mainly lives well below the tidal zone in algae on rocks. Habitats—Known only from among algae in shallow water (G. pisinna) or in sand (G. sulci- fera); for details see under these species. G. pis- inna may live as deeply as 8 fathoms; a probably adventitious shell of G. sulcifera came from 287 fathoms. Larval ecology—The fairly small range of varia- tion in the diameter of the first whorls of the six known taxa of Gabrielona (0.20-0.29 mm.) and the small sizes presumably indicate that the full- grown larval shells are small and relatively uni- form in size, and that the larvae are all pelagic and planktotrophic. Abnormalities—A_ striking series of abnormal- ities possibly caused by an individual living in an unusual habitat is described under G. sulcifera. Abnormal growth caused by incrustations are rare in Gabrielona; one case is reported under G. nepeanensis. [04-154] May 30, 1973 Synonymy- 1917 Gabrielona Iredale, Proc. Malac. Soc. London 12:322 [listed], 327. Type-species (by monotypy): Phasianella nepeanensis Gatliff and Gabriel, 1908.—1929, Thiele, Handb. syst. Weichtierkunde, Jena, 1:70.—1938, Wenz, Handb. Palaozool., Berlin, 6(1)Teil 2 [Prosobranchia], p. 362,— 1958, Robertson, Johnsonia 3(37):246-260. Excluded species—In 1958 (pp. 253, 257), I sug- gested that three American Miocene (or Plio- Pleistocene)species might belong in Gabrielona: Tricolia (Eulithidium) hadra Woodring, Didia- nema P waltonia Gardner, and Tricolia P syn- toma Woodring. T. hadra is here referred defi- nitely to Gabrielona, but subsequent study of the holotypes of the other two species has shown that neither belongs in the Phasianellidae. Two upper Tertiary species from northern Venezuela described as Gabrielona are treated under Ex- cluded Species on p. 61. The possible second Western Atlantic Recent species of Gabrielona (Robertson, 1958, p. 259), from Brasil, proves upon restudy to be a depau- perate, low-spired Tricolia. Gabrielona nepeanensis (Gatliff and Gabriel, 1908) (Pls. 36-39) Range— Recent: known only from South Austra- lia and Victoria, Australia. Possibly occurs also at Tasmania, but not yet known there. Chief distinguishing characters—The shell dif- fers from those of all other known species in the genus except G. pisinna in having both columel- lar denticles (ed; and ed,), in lacking axial or spiral sculpture, and in opercular characters (see under G. raunana and G. sulcifera for differ- ences). Differs from all other species, including G. pisinna, by its complex and varying but con- sistently distinct color patterns (the adults always with a colorless spiral band opposite the palatal sulcus). For detailed differences from G. pisinna, the most similar species, see under that species. Abundance—36 shells from beach sand avail- able, 1 with operculum in place in aperture; none live-collected (probably lives below the tidal zone). Abnormal shell—One shell (PI. 36, fig. 6) has a double outer lip near the suture, a columellar callus slightly detached from the palatal area, and the upper columellar denticle (cd,) fainter than usual. Some incrustation must have inter- fered with normal growth. INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 Gabriclona 45 Shell description—Attains length of 1.9 mm., width of 1.7 mm., and 3.9 whorls; spire angle 95°-105°; length invariably equals or exceeds width (except for small shells); fairly thick but slightly translucent to fairly transparent. Proto- conch insert, slightly inflated, smooth, white, not demarcated from teleoconch. First whorl 0.24- 0.28 mm. in diameter. Penultimate and _ last whorls: slight flattening below suture but no dis- tinct shoulder (PI. 38, fig. 1); surface smooth ex- cept for axial growth lines and slight wrinkles, fairly shiny. Colors: pale to dark pink (rarely, tinged with orange), and white. Patterns: alter- nating pink and white subsutural marks, each becoming divided by a virtually colorless spiral band directly opposite the palatal sulcus; 9-12 paired pink marks on last whorl; predominant below subsutural area: axially aligned wavy pink stripes or irregular marks; subperipheral series fairly faint pink and white marks; inner umbilical area colorless, surrounded by spiral series short white axial stripes (commonly, partially coa- lesced); rarely, zigzag pale pink ‘stripes or irregu- lar marks entirely replace usual pattern (PI. 36, figs. 3-4). Outer lip and callus on upper parietal area fairly thin; no palatal denticle; palatal sul- cus high in aperture; both columellar denticles present, most prominent on large shells (PI. 38, fig. 2). Columellar lip thickened adjacent to cen- tral part umbilical channel, and an escarpment present; escarpment to left of umbilical channel fairly faint to prominent, arising fairly low off the outer edge of columellar lip; umbilicus nar- row to fairly wide. Shell measurements (mm.)— length width no. whorls 1.92 1.66 3.9 large; South Australia 1.56 1.46 3.3 average; South Australia 0.80 0.87 2D smallest; South Australia Plate 37. Gabrielona nepeanensis (Gathff and Gabriel). Vic- toria, Australia. Original figures of holotype (from Gatliff and Gabriel, 1908, pl. 21, figs. 9-10), enlarged. Both x20. Fig. | incorrectly shows a prominent shoulder (compare PI. 36, fig. 10). [04-155] 46 Gabrielona Plate 38. Gabrielona nepeanensis (Gatliff and Gabriel). Fig. 1. Outline of large shell, x17. Fig. 2. Aperture, x33; ar, aper- tural ridge; ed, and edz, upper (outer) and lower (inner) col- umellar denticles (respectively); ps, palatal sulcus; uc, umbil- ical channel. Figs. 3-5. Operculum. Internal surface, longitu- dinal section, and external surface (respectively). All x33. Operculum-(Pl. 38, figs. 3-5). Only a single, abraded operculum of this species has been available, and it has been difficult to determine the sculpture of its external surface. The crest of the spiral ridge near the edge seems to be round- ed, and on the central region—between the outer, spiral area and the central area—there is a low, spiral ridge extending to a prominence on the non-spiral columellar margin. The outline of the columellar margin at the junction of the spiral and non-spiral sectors is shallowly concave. Otherwise the operculum of G. nepeanensis seems to be like that of G. pisinna, only larger; both are fairly thin. Synonymy— 1908 Phasianella nepeanensis Gatliff and Gabriel, Proc. Roy. Soc. Victoria, n.s., 21(1): 366, pl. 21, figs. 9-10 [re- produced here, PI. 37] (Flinders, Western Port; Ocean Beach, near Point Nepean [both Victoria, Australia]). —1917, Iredale, Proc. Malac. Soc. London 12(6): 322 [listed], 327 [the only named species included in Gab- rielona but not formally transferred]. 1938 Gabrielona nepeanensis (Gatliff and Gabriel). Cotton and Godfrey, Malac. Soc. South Australia Publ. 1: 9; 1945, Cotton, Trans. Roy Soc. South Australia 69(1): 165; 1958, Robertson, Johnsonia 3(37): 257, pl. 137, figs. 2-3, pl. 140, fig. 1; 1959, Cotton, South Australian Mollusca, Archaeogastropoda, Adelaide, pp. 270-271, fig. 185 [shell shape highly inaccurate], p. 347 [listed]. Robert Robertson Phasianellidae Types—The holotype of Phasianella nepeanensis Gatliff and Gabriel (Pl. 36, fig. 10; Pl. 37) is now at the National Museum of Victoria (no. F543), Melbourne, Australia. A small paratype is at the Australian Museum (no. C.45057), Sydney (Rob- ertson, 1958, pl. 140, fig. 1). Gatliff and Gabriel did not mention the number of specimens avail- able to them, and recorded the species from two localities 23. miles apart. Cotton and Godfrey (1938) selected “Flinders, Victoria” as the type- locality. However, in 1945 and_ subsequently, Cotton has stated that the type-locality is “near Point Nepean,” whence (judging by the specific name) the holotype came. Locality records (see map, Pl. 39; literature and uncertain records circled—SOUTH AUSTRALIA: Port Lincoln (“appar- ently nepeanensis,” Iredale, 1917; Cotton, 1945); Mouth of Middle River, N. coast Kangaroo I. (from beach sand, 1954, B. Daily, via M. F. Glaessner and G. L. Harrington); Port Adelaide (Calvert Coll., both ANSP); Robe (Cotton, 1945). VICTORIA: Port Fairy (in beach sand, 1923, H. A. Pilsbry, ANSP); ocean beach near Point Nepean [38 mi. S.S.W. of Melbourne] (Gatliff and Gabriel, 1908; Natl. Mus. Vict.: Austral. Mus.); Flinders [47 mi S. of Melbourne], Western Port (Gatliff and Gabriel, 1908). All three of the circled locality records in South Australia are questionable because Cotton (1945, 1959) mentioned having difficulty distinguishing G. nepeanensis from “Pellax virgo,” and the locality data with all the available specimens ex- cept one juvenile shell from Kangaroo Island seem not wholly reliable either. A large series (30 shells) is from the Calvert Collection (col- lector not recorded), labeled “Pt. Adelaide” is suspect because this is the largest port in the area. Gabrielona pisinna Robertson, new species (Pls. 39-42) Range—Recent: known only from Mauritius, In- dian Ocean, and New Caledonia, eastern Melan- esia. Perhaps widespread in the tropical Indo- Pacific around high islands. Chief distinguishing characters—The shell, usual- ly less than 1 mm. in length, is full-grown at a smaller size than in any other known phasianel- lid. It resembles young G. nepeanensis in size and number of whorls. (Further resemblances: has both columellar denticles, an unsculptured sur- face, and a similar operculum.) That G. pisinna is full-grown at a smaller size than G. nepean- ensis is shown by the complete development on the larger shells of the lower columellar denticle (ed,). Further differences from G. nepeanensis: first whorl smaller; spire lower (except for some [04-156] May 30, 1973 MALDIVE 4 © is. CHAGOS __ is rata) €0C06 - KEELING ATOLLS : | Plate 39. Geographical distributions and records of Gabriel- ona nepeanensis (Gatliff and Gabriel), Gabrielona species? (Iredale, 1917), and Gabrielona pisinna Robertson, which large shells, the width of G. pisinna invariably exceeds the length); aperture smaller than in young G. nepeanensis of comparable size (com- pare Pl. 40 with Pl. 36, fig. 1); color patterns dif- ferent (finer), and reddish coloration darker. Relationships—A dwarf, tropical homologue of G. nepeanensis, which might need to be ranked as a subspecies if the form occurring at Lord Howe Island (map, Pl. 39) is intermediate (see p. 44). Abundance—68 specimens available: 2 empty shells from Mauritius (1 with operculum), and 66 specimens from New Caledonia (10 with oper- cula, 7 of these live-collected). Habitat—All 7 of the live-collected specimens came from washings from algae collected in 0-3 ft. on Récif Ricaudy, New Caledonia, on rocks near shore on the fringing reef. An empty but fresh shell with 4 drill holes and an operculum in place in the aperture came from a depth of 8 fathoms at Mauritius (see under Locality rec- ords). (Many of the shells have these drill holes.) Shell description—Attains length of 1.1 mm., width of 1.1 mm., and 3.0 whorls; spire angle 115°-125°; width exceeds length (excepting some large shells); fairly thin and translucent to transparent. Protoconch insert, slightly inflated, smooth, white, not demarcated from teleoconch. First whorl 0.20-0.24 mm. in diameter. Penulti- mate and last whorls: slight flattening below su- ture but no distinct shoulder (Pl. 41, figs. 2-3); surface smooth except for axial growth lines and slight wrinkles, shiny; fine spiral sulci on base of small shells (Pl. 41; fig. 1). Colors: dark pink to bright red (rarely, tinged with amber [faded?]), and white. Patterns: white spiral band below the suture, coalescing with variably-shaped white subsutural patches (7-9 on last whorl), lower INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 Gabrielona 47 *) MARIANA Nokes * Guam : Sopon « oO cof NEW ZEALAND 1 160° 160° G, NEPEANENSIS perhaps is widespread in the tropical Indo-Pacific but which presently is known only from the two areas 7300 miles apart. edges of which are slightly below or directly op- posite the palatal sulcus; pink or red marks or non-axial stripes alternate with white subsutural patches; predominant below subsutural area: fairly regular axial or very steeply ascending dark stripes; subperipheral series of white marks alternating with non-axial dark stripes or axially paired marks; umbilical area closely surrounded by spiral series of short white axial stripes (fair- ly commonly, partially coalesced); very rarely, subsutural patches and subperipheral white marks coalesce into irregular axial white bands (Pl. 40, fig. 2). The aperture averages relatively Plate 40. Gabrielona pisinna Robertson. Fig. 1. Mauritius. Figs. 2-6. New Caledonia. Figs. 4-6. Holotype. All x30. [04-157] 48 Gabrielona Robert Robertson Phasianellidae Plate 41. Gabrielona pisinna Robertson. Fig. 1. Smallest spec- imen, x60. Figs 2-3. Outlines of largest shells, both x33. Fig. 2. Unusually high-spired shell. Fig. 4. Aperture, x60; ar, aper- tural ridge; ed, and cd, upper (outer) and lower (inner) col- umellar denticles (respectively); ps, palatal sulcus; uc, um- bilical channel. Figs. 5-7. Operculum. Internal surface, longi- tudinal section, and external surface (respectively); ¢, lower edge of callus. All x60. slightly wider than in G. nepeanensis. Outer lip and callus on upper parietal area thin; no palatal denticle; palatal sulcus high in aperture, slightly variable in position (commonly, fairly faint); both columellar denticles present, most prominent on large shells (Pl. 41, fig. 4). Columellar lip slightly thickened adjacent to central part umbilical channel, and an escarpment, when present, faint; escarpment to left of umbilical channel fairly prominent, arising fairly low off outer edge col- umellar lip; umbilicus fairly wide. Plate 42. Gabrielona pisinna Robertson. Half of one trans- verse row of radular teeth, showing (from left to right) the 5-cusped central (or pseudocentral?), the three laterals on the right side, and the entire row of marginals. The cusps of the central and laterals are stippled. New Caledonia. x1050. Shell measurements (mm. )— length width no. whorls 1.09 1.10 3.0 largest; New Caledonia 0.91 0.93 2.8 holotype; New Caledonia 0.70 0.75 2.6 average; Mauritius 0.34 0.42 1.7 smallest (Pl. 41, fig. 1); New Caledonia Operculum—(PI. 41, figs. 5-7)- The spiral ridge near the edge on the external surface of the fair- ly thin operculum is sharply crested except near the columellar margin where it is rounded. The inner margin of the ridge is angled or rounded. The central region has an outer, spiral area with a mat surface contrasting with a shiny, more transparent central area. The single operculum available from Mauritius differs from those from New Caledonia in having a faint spiral ridge be- tween the two areas (like that of G. nepeanen- sis). Fairly prominent wrinkles parallel to the non-spiral columellar margin are present to the left of the central part of the callus. The lower margin of the callus (c) varies in position. Radula (Pl. 42; 2 studied)—Attains length of 0.37 mm., width of 0.08 mm., and with as many as 20 very strongly curved transverse rows of teeth (including a few nascent rows). Central (or pseudocentral?) 5-cusped, laterally winged ante- riorly, and with posterolateral projections on the base. Three laterals, each with long pointed cusps (the two innermost generally with 5 cusps, and the outermost with 6); the two outermost laterals have the largest distal portions, and the outer lateral is positioned posterior (rather than lateral) to the morphologically middle lateral; the [04-158] May 30, 1973 base of the innermost lateral bears a posterolat- eral peg that fits into the middle of the inner edge of the base of the middle lateral, and the two outermost laterals have large bases that ex- tend posterolaterally beneath the marginals. As many as 13 pairs of marginals (lowest count 10), each row widely overlapping the row posteriad. All the marginals have elongate distal portions, and the innermost of these are serrate on the outer (posterior) edge; the serrations become fin- er outwards and are absent altogether on the small outermost teeth. Types—The holotype (Pl. 40, figs. 4-6), from Récif Ricaudy, near Noumea, New Caledonia, is at the Academy of Natural Sciences of Philadel- phia (no. 301611). So also are paratypes from the type-locality (no. 271062) and from other lo- calities: Récif de Gatope, New Caledonia (nos. 267567 and 267568), and Mauritius (nos. 273188 and 273328). Paratypes from New Caledonia will be distributed to USNM, MCZ, BM, and IrSnB (2 shells each). Derivation of new name—Latin, pisinnus, little. Locality records (see map, Pl. 39)—MAURITIUS: % mi. N.N.E. of Flic en Flacq Pt. (1-10 ft., from Caulerpa wash- ings, Nov., R.E.M. Ostheimer & V. Orr Maes, Sta. M 203): Black River Bay, 1 mi. W.N.W. of mouth Black River, both W. coast (dredged 8 fms. [1 empty but fresh shell], coarse sand, broken shell, very little weed, Nov. 5, both 1960, R.E.M. Ostheimer, J. de B. Baissac & V. Orr Maes, Sta. M 208, both ANSP). NEW CALEDONIA: Grand Reéif de Ga- tope, 7% mi. W. of Voh (dredged 6-18 ft., inner edge of bar- rier reef, sand, weed, coral rubble, Dec. 31, 1960 & Jan. 2, 1961, Stas. K 538 & K 539); E. end Reécif Ricaudy, 2% mi. S.S.E. of Noumea (0-3 ft., Jan. 11, 1961, both G. & M. Kline & V. Orr Maes, Sta. K 553, both ANSP). Gabrielona raunana Ladd, 1966 (Pls. 43-50) Range—Recent: known only from Eniwetok (an atoll), northwestern Marshall Islands (G. raunana raunana Ladd), and from the Loyalty Islands and New Caledonia (all high islands), eastern Melan- esia (G. raunana goubini Robertson). Chief distinguishing characters—This is the only species in the Phasianellidae having a shell with prominent axial sculpture. The plicae are most prominent and regularly arranged on the second whorl, and are obscure or absent at and near the periphery of the last whorl of large shells. Also differs from other species in the genus as fol- lows: spiral keel on protoconch; all apertural denticles lacking; operculum with obliquely radi- al sulci on part of external surface, and with INDO-PACIFIC MOLLUSCA vol. 3, no. 14 Gabrielona 49 structural radial lamellae at and near the spiral outer edge. The outlines of the shells and color patterns of both subspecies are distinct from each other and from those of the other species. Remarks—No live-collected specimens are avail- able of either subspecies; most of the shells are beach worn or subfossil. G. raunana, known only from populations in two areas slightly more than 2,000 miles apart, seems thus to be a relict spe- cies. Shell descriptions, Types, Locality records, etc. See under G. raunana raunana and G. raunana goubini. Operculum (Pl. 44)—Only 2 opercula of G. raunana raunana and 8 of G. raunana goubini are available, and because all these are abraded or corroded—making description and illustration of the original sculpture difficult—and because it is doubtful whether the seeming slight differ- ences between the two subspecies are real, they are discussed together here. The operculum in best condition is a small one from G. raunana raunana. Relative to those of G. nepeanensis and G. pisinna, the operculum of G. raunana (both sub- species) is thick and comprises fractionally more whorls (comparing opercula of the same size). The external surface has a central region with an outer, spiral area distinctive in having prominent, obliquely radial sulci (irregular and varying in spacing), and a central, smoothish area with a fairly thick callus. The crest of the spiral ridge is Plate 43. Gabrielona raunana Ladd. Thinly coated with mag- nesium oxide to accentuate the sculpture and obscure the color pattern. Figs. 1-2. G. raunana raunana Ladd. Eniwetok, Marshall Islands. Figs. 3-4. G. raunana goubini Robertson. Lifou, Loyalty Islands. All x20. [04-159] 50 Gabriclona Phasianellidae Robert Robertson near the outer edge, and (at least on G. raun- ana raunana) its whole surface is irregularly wrinkled and knobbed, the wrinkles tending to be aligned with the adjacent obliquely radial sulci. The erosion of the external surface makes sev- eral features obscure. The spiral ridge is strongly abraded on all the opercula available from G. raunana goubini. On all but one of these there is a deep but irregular central pit that may be an erosional feature; this is margined by an irregu- lar but steep escarpment at the lower margin of the callus (c). On the unpitted operculum of G. raunana goubini, two spiral sulci are near the center and there is no steep escarpment border- ing the callus; this perhaps is the uneroded orig- inal sculpture. Seen in transparency, near the outer edge of the operculum are structural radial lamellae at right angles to the horizontal plane of the oper- culum. These are closely and regularly spaced and extend a uniform distance from the edge. Most prominent at the edge of the operculum of G. raunana raunana, these project from the sur- face as external lamellae, thus causing the out- line to be finely notched. These external lamellae perhaps are erosional features. Gabrielona raunana raunana Ladd, 1966 (Pl. 43, figs. 1-2; Pl. 44, figs. 4-6; Pls. 45-47) Range—Recent: known only from subsurface de- posits on Eniwetok Atoll, northwestern Marshall Islands (obtained from drillings). Perhaps wide- spread at atolls in Micronesia or the whole tropi- cal northwest Pacific. Chief distinguishing characters—The shell of this subspecies has fairly regularly spaced but commonly coalesced colorless or white spots, each one surrounded by 6 others. More differ- ences are given under G. raunana goubini. Sculp- ture is almost identical in the two subspecies. Abundance—32 subfossil shells available, 1 with an operculum in place in the aperture; 1 loose operculum. Shell description—Attains length of 2.1 mm., width of 2.1 mm., and 3.4 whorls; spire angle 110°-125°; width invariably equals or exceeds length; fairly thick to fairly thin and opaque to fairly transparent. Protoconch insert, smooth ex- cept for slightly descending spiral keel, white, slightly demarcated from teleoconch, 0.9 whorl. First whorl 0.26-0.29 mm. in diameter. Penulti- mate and last whorls: flattening below suture, and faint shoulder (Pl. 46, fig. 1). Sculpture of second whorl: 18-30 strong axial plicae, convex towards outer lip, highest, most sharply crested and most widely spaced on first half where a fairly prominent to fairly obscure spiral cord extends from the keel on protoconch. On later whorls pli- cae smaller, crests rounded, more closely spaced, and less regular in arrangement and _ structure, commonly with intercalated secondary plicae be- low suture and with divarications and anasto- moses below periphery. A faint spiral cord sur- rounds the umbilical area of young shells; rarely, faint reticulations on base of medium-sized shells. On last whorl irregular axial plicae most promin- ent below suture and on base, absent near peri- phery of large shells where surface is smooth except for axial growth lines. Surface shiny, Colors: pale (faded?) to fairly pale pink or yel- lowish brown, and white. Patterns: more or less quadrate pink or brownish subsutural patches (6-10 on last whorl), lower edges directly opposite palatal sulcus; surface almost entirely covered with fairly regularly spaced spots, each one sur- Plate 44. Gabrielona raunana Ladd. Opercula. Figs. 1-3. G. raunana goubini Robertson, x33. Figs. 4-6. G. raunana raun- ana, x60. Internal surface, longitudinal section, and external surface of each; c, lower edge of callus. [04-160] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 Gabrielona 5] Plate 45. Gabrielona raunana raunana Ladd. Subsurface Re- cent deposits, Eniwetok Atoll, Marshall Islands. Fig. 7. Holotype. Figs. 1-2, x40; Figs. 3-7, x20. rounded by 6 others; spots white in pale areas alternating with subsutural patches and colorless elsewhere except (somewhat commonly) for sub- peripheral spiral series and (rarely) spiral series on base; commonly: spots irregularly coalesced; ground color pale; fairly commonly: subperipheral spiral series irregular, slightly darkened markings alternating with white-spotted areas; closely sur- rounding umbilicus: spots variously coalesced in- to irregular, steeply descending white stripes on colorless ground; fairly rarely: wavy, axial dark- ened bands extending from subsutural patches to base; rarely: almost a uniform pink with no spots near periphery. Outer lip and callus on upper parietal area somewhat thin to fairly thick; no palatal denticle; palatal sulcus high in aperture, Plate 46. Gabrielona raunana raunana Ladd. Fig. 1. Outline of large shell, x17. Fig. 2. Aperture, x33; ar, apertural ridge, ps, palatal sulcus; uc, umbilical channel. neither columellar denticle present (Pl. 46, fig. 2). Columellar lip not thickened, and steep slope into umbilical channel but no escarpment; es- carpment to left of umbilical channel prominent, arising quite high off the outer edge of the col- umellar lip; umbilicus fairly narrow to wide (PI. 46, fig. 2). Shell measurements (mm.)— length width no. whorls 2.09 2.09 3.4 largest 1.60 1.67 3.1 average; holotype 0.89 1.07 2.4 smallest O perculum—See under G. raunana (species). Synonymy— 1966 Gabrielona raunana Ladd, [U.S.] Geol. Surv. Prof. Paper 531, pp. 13 & 17 [listed], 54 [described], pl. 10, figs. 1-5 (Recent, Eniwetok Atoll). Types—The holotype (Pl. 45, fig. 7), out of a drilling from 20-45 ft. below land surface Eluge- lab, Eniwetok, is at the United States National Museum (no. 648319), Washington, D.C. So also are all the paratypes except 3 donated to the Academy of Natural Sciences of Philadelphia (no. 302131). Locality records (see map, Pl 47)— MARSHALL ISLANDS: Elugelab (20-60 ft. deep), Parry (30-45 ft., 90-110 ft., 1865- 1895 ft.), and Mujinkarikku (35-40% ft.), all Eniwetok Atoll (from 8 drill holes in Recent subsurface deposits; 30 shells and 1 loose operculum at depths of 20-60 ft., 1 shell from 90-110 ft., and 1 probably adventitious shell in lower Mio- sene strata at 1865-1895 ft., all about 1952, IL.S. Ladd, USNM & ANSP). For an account of the drilling operations which yielded most of the specimens, see Ladd and Schlanger (1960, [U.S.] Geol. Surv. Prof. Paper 260-Y, pp. i-iv, $63- 905). [04-161] 52. Gabrielona | g 4° ' Iwo Jima . Mores poo SW rormosA Cs “MARIANA 2 Is. G. RAUNANA RAUNANA ye ye o go" Enwetdh@ \ 7 © MARSHALL IS. |, Oth Sasle ‘ “Y = + +, Wot/e AO oe Sa Cee ee Ponape +, Majuro ROLINE Me Se C Ss, 4 . | See oo us eae a 5 | New NF, ee ef ) CALEDONIA WP a 1 G. RAUNANA GOUBINI 20° \40° Plate 47. Geographical records of Gabrielona raunana_ rau- nana Ladd and Gabrielona raunana goubini Robertson. Robert Robertson Phasianellidae Gabrielona raunana goubini Robertson, new subspecies (Pl. 43, figs. 3-4; Pl. 44, figs. 1-3; Pls. 47-50) Range—Recent: known only from Lifou, Loyalty Islands, and Ile des Pins, New Caledonia, eastern Melanesia. Perhaps widespread around high is- lands in Melanesia or the whole tropical south- west Pacific. Chief distinguishing characters—The shell dif- fers from that of the nominate subspecies as fol- lows: spire averages higher and aperture rela- tively smaller; first whorl averages smaller; pli- cae finer on medium-sized shells; color pattern almost invariably with steeply descending pink or yellowish brown stripes, and white subsutural patches commonly ring-shaped. Abundance—1,116 shells available, 8 with an operculum in place in the aperture. All the speci- mens were sorted from beach sand; some are freshly dead, but many are worn or broken and some are bleached. Plate 48. Gabrielona raunana goubini Robertson. Lifou, Loyalty Islands. Fig. 7. Holotype. Figs. 1-2, x40; Figs. 3-10, x20. [04-162] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 Gabriclona — 53 Remarks—In view of the apparent abundance of this subspecies in beach sand it is remarkable that it has not been named previously, in spite of rather extensive study of small marine shells from the Loyalty Islands (Tomlin, 1936, Proc. Malac. Soc. London 22(3): 145-152). Color variation—Most of the shells have pink markings; an infrequent color form is entirely de- void of pink and has pale yellowish brown mark- ings; intermediates have the pink followed by the yellowish markings. Only a few shells were not readily sorted into one of these three categories. Of the 1,088 shells from Lifou, approximately 938 (86%) are the pink form, 112 (10%) are inter- mediates, and 38 (4%) are the yellowish form. The frequencies of the three color forms are comparable in the much smaller sample (28 shells) from Ile des Pins: 18 (64%) pink, 7 (25%) intermediates, and 3 (11%) yellowish. Shell description—Attains length of 2.4 mm., width of 2.2 mm., and 3.9 whorls; spire angle 90°-115°; width fairly commonly exceeds length; fairly thick but slightly translucent to fairly trans- parent. Protoconch like that of G. raunana raun- ana. First whorl 0.23-0.28 mm. in diameter. Sub- sequent whorls: flattening below suture, and faint to obscure shoulder (PI. 50, fig. 1). Sculp- ture like that of G. rauwnana raunana except pli- cae (PI. 50, fig. 2) finer, spiral cord on first quad- rant(only) of second whorl faint or absent, and no reticulations on base of medium-sized shells. Surface shiny. Colors: fairly pale to dark pink, pale yellowish brown, and white. Patterns: vari- ably-shaped white subsutural patches (7-9 on last whorl), commonly ring-shaped with axial stripe extending to suture, lower edges near or direct- ly opposite palatal sulcus; white subsutural patches alternating with variable yellowish brown or pinkish marks; predominant below sub- sutural area: steeply descending pink and/or yel- lowish brown stripes; commonly: subperipheral series small, irregular white marks alternating with slightly darkened pinkish or yellowish brown marks; fairly commonly: similar series small white marks on base, with darkened pink- ish marks; closely surrounding umbilicus: steeply descending, partially coalesced white — stripes: very rarely: all descending stripes zigzag or bro- ken into irregular marks (PI. 49). Aperture aver- ages slightly narrower and _ relatively smaller than that of G. raunana raunana; outer lip some- what thick; callus on upper parietal area thin to fairly thick; apertural denticles, palatal sulcus, columella and umbilical area as in G. raunana raunana, except escarpment to left of umbilical channel fairly faint to prominent and umbilicus narrow to quite wide. Shell measurements (mm.)— length width no. whorls 2.42 2.16 3.9 largest 1.92 1.80 3.5 average; holotype 0.90 1.01 2.6 smallest Plate 49. Gabrielona raunana goubini Robertson, Rare color patterns, Figs. 1-3. Lifou, Loyalty Islands. Fig. 4. Tle des Pins, New Caledonia. All x20. Plate 50. Gabrielona raunana goubini Robertson. Fig. 1. Out- line of large shell, x17. Fig. 2. Apex, x33; p, protoconch. Fig 3. Aperture, x33; ar, apertural ridge; ps, palatal sulcus; uc, umbilical channel. [04-163] 54. Gabrielona Operculum—See under G. raunana (species). Types—The holotype (PI. 48, fig. 7) from Lifou, Loyalty Islands, is in the Dautzenberg Collection at the Institut royal des Sciences naturelles de Belgique, Brussels. So also are 1,102 paratypes from Lifou and Ile des Pins. Ten more paratypes from Lifou are retained at the Academy of Nat- ural Sciences of Philadelphia (no. 302624), and 3 from Lifou have long remained unidentified at the United States National Museum _ (no. 422601), Washington, D. C. Derivation of new name—Named for Goubin, the collector who meticulously sorted out from beach sand 1,085 of the shells. Locality records (see map, Pl. 47)—. LOYALTY ISLANDS: Ile Lifou (Goubin, IrSnB & ANSP; Moss, USNM). NEW CAL- EDONIA: Ile des Pins (Lambert, IrSnB). Robert Robertson Phasianellidae Gabrielona hadra (Woodring, 1928) (Pls. 51-53) Range—Middle Miocene or Plio-Pleistocene: known only from the Bowden Formation, south- eastern Jamaica, Greater Antilles. Presumably was widespread in the Caribbean area. (On ap- parent endemism in the Bowden Formation, see W. P. Woodring, 1965, Science 148 (3672): 961- 963.) Chief distinguishing characters—The shell at- tained a larger size than that of any Recent spe- cies in the genus. G. hadra is distinct also in hav- ing a deeply embayed columellar lip below the junction with the palatal wall. Otherwise, G. hadra closely resembles G. sulcifera (the sculp- ture of the second whorl is similar, the palatal Plate 51. Gabrielona hadra (Woodring). Middle Miocene (or Plio-Pleistocene), Bowden Formation, Jamaica. Fig. 4. Larg- est known specimen of any Gabrielona (surface eroded). Figs. 5-7. Holotype. All x20. [04-164] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 ul on Gabrielona Plate 52. Gabrielona hadra (Woodring). Figs. 1-2. Outlines of shells, showing variation, both x17. Fig. 2. Largest known specimen of any Gabrielona (surface eroded). Figs. 3-4. Aper- tures, showing variation; both x33; ar, apertural ridge; cdo, lower (inner) columellar denticle; ps, palatal sulcus; uc, um- bilical channel. Figs. 1 and 4. Holotype. sulcus is near the middle of the aperture, and on- ly the lower columellar denticle (edz) is present). Additional differences from G. sulcifera: first whorl averages smaller; spiral sculpture on sec- ond whorl less prominent, disappearing before the beginning of the third whorl; no spiral sulci on later whorls; color patterns different. Abundance—21 shells currently available, none with opercula (36 paratypes not available; see under Types). Remarks—This is the only known fossil species undoubtedly belonging in the genus, and clearly is a precursor of the Recent Caribbean G. sulci- fera. The relative abundance of G. hadra in the Bowden Formation is noteworthy in view of the rarity of its living descendant. The surface of the shell is most resistant to cor- rosion where there were white markings, which on some shells are preserved as low projections. Shell description—Attained length of 3.3 mm., width of 3.1 mm., and 4.6 whorls; spire angle 95°-120°; length invariably exceeds width (ex- cept possibly for small shells), but outline vari- able; fairly thick to thick, and opaque (fresh shells might be translucent or transparent). Pro- toconch slightly exsert, slightly inflated, smooth, whitish, slightly demarcated from teleoconch, 1.1-1.2 whorls. First whorl 0.23-0.25 mm. in di- ameter. Penultimate and last whorls: slight flat- tening below suture and faint shoulder (PI. 52, figs. 1-2). Sculpture of second whorl: 3 fairly low spiral keels on first quadrant following proto- conch, each about equal in prominence; keels gradually becoming spiral cords, commonly with 2 more cords intercalated; very fine axial threads; all sculpture gradually disappearing on third or fourth quadrant, commonly with no spi- ral sulci; on later whorls surface wholly smooth except for fine axial growth lines. Surface shiny. Colors: pale (faded?) reddish or purplish brown, and white. Patterns: irregularly shaped whitish subsutural patches (8-11 on last whorl), alternat- ing with brownish areas; lower edges of patches irregular, not correlated with position of palatal sulcus; predominant below poorly demarcated subsutural area: ground color pale reddish brown; 2-3 spiral series irregular white markings (commonly crescentic, concave towards outer lip) that are smaller than the subsutural patches; 2 of these series near (above and below) periphery (upper one fairly uncommonly absent), and third on base (6-9 markings); umbilical area com- monly tinged with purple. Aperture shape like [04-165] 56 Gabrielona that of G. sulcifera; outer lip fairly thin to thick; callus on upper parietal area thin to thick; pala- tal denticle absent; apertural ridge (ar) shallow to fairly deep in aperture (Pl. 52, fig. 3); pala- tal sulcus near middle of aperture; upper colu- mellar denticle (cd,) absent; lower columellar denticle (cd,) present, prominent or low and wide (Pl. 52, figs. 3-4). Columellar lip not thick- ened, deeply embayed below junction with pala- tal wall; fairly shallow slope into wide umbili- cal channel; escarpment to left of umbilical channel prominent, arising fairly high to high off outer edge columellar lip, and commonly the right-hand edge of a ridge; umbilicus wide. Shell measurements (mm.)— length width no. whorls 3.30 3.06 4.6 largest 2.01 2.44 4.2 average; holotype 2.10 2.04 4.0 fairly small Synonymy— 1928 Tricolia (Eulithidium) hadra Woodring, Carnegie Instit. Washington Publ. 385 (Miocene Mollusks from Bow- den, Jamaica; Part I), pp. 16 [name listed], 420-421, pl. 34, figs. 10-11.—1958, Robertson, Johnsonia 3(37): 253, 257 [provisionally referred to Gabrielona]. Types—The holotype of Tricolia (Eulithidium) hadra Woodring (PI. 51, figs. 5-7), from near Bowden, Jamaica, is at the United States Nation- al Museum (no. 369556), Washington, D.C. This presumably was one of the “37 specimens in the Duerden Collection” mentioned by Woodring. Plate 53. Geographical records of Gabrielona sulcifera Rob- ertson (Recent) [round spots] and Gabrielona hadra Robert Robertson is dl @ a ry a | | ‘Sia sticrox " @ANTIGUA | TAMAR EM Mio | ----ae eben ooo “pe | eS aa | v \—'5 y | = | =I a | | , \s CARIBBEAN | 2 \s . SPD Phasianellidae The remaining 36, which can be considered para- types, were not found in the paleontological col- lection from Johns Hopkins University on deposit at USNM (May, 1965). Twenty topotypes from the Henderson collection are at USNM (no. 135509). Fossil record (see map, Pl. 53, black triangle)— JAMAICA: near Bowden, St. Thomas Parish (1894, J.B. Henderson, Jr.; 1899, J.E. Duerden; in thin bed imperfectly consolidated gravel in a marly matrix; USNM). Gabrielona sulcifera Robertson, new species (Pls. 53-57) Range—Recent: known only from off northwest- ern Cuba, Greater Antilles, from the Virgin Is- lands, and from Antigua, Lesser Antilles. Pre- sumably widespread around high _ islands throughout the West Indies, but very rarely col- lected. Chief distinguishing characters—This is the only species in the genus having a shell with promin- ent spiral sculpture. This begins on the second whorl as keels, which soon become cords. On later whorls the cords are reduced and_ broad- ened to interspaces between sulci. These sulci are absent at and near the periphery of the last whorl of large shells. Also differs from other Re- cent species in the genus as follows: very fine ax- ial threads on second whorl; color pattern with yt be Boe oly te G. HADRA|™ OM. Mio.) “a (Woodring), its Middle Miocene (or Plio-Pleistocene) pre- cursor [triangle]. [04-166] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 on ~l Gabrielona Plate 54. Gabrielona sulcifera Robertson. Antigua, Lesser Antilles. Fig. 4. Holotype. All x20. no demarcated subsutural area; palatal sulcus near middle of aperture and not correlated with color pattern on external surface; palatal denticle can be present; only lower columellar denticle (ed) present (abnormally, slight trace upper col- umellar denticle); operculum with prominent spi- ral cord at inner edge marginal spiral ridge. For differences from G. hadra, see under that species. Relationships—Judging by the presence and kind of sculpture (even though spiral and not axial), the fairly thick, generally similarly-shaped shells, and the two opercular resemblances, G. sulcifera seems more closely related to G. rau- nana than to the other two Recent species. How- ever, the differences are far greater than those between G. nepeanensis and G. pisinna. Abundance—14 specimens available, only 1 live- collected and with an operculum. Habitat—The single live-collected specimen and 11 empty shells were all dredged together at an unrecorded depth at Antigua, and were in “deep, fine sand and shells.” To have been in or near English Harbour, the depth must have been less than about 20 fathoms (H.O. chart 366). The empty, drilled, abnormal shell from 287 fathoms off northwestern Cuba probably was adventi- tious at that depth. Shell description [see also section on ab- normal shell|—Attains length of 2.4 mm., width of 2.3 mm., and 4.3 whorls; spire angle 90°- 105°; length exceeds width, and even small shells high-spired; outline fairly variable (Pl. 56, figs. 1-3); fairly thick and slightly translucent to slightly transparent. Protoconch slightly exsert, slightly inflated, smooth, white, fairly prominent- ly demarcated from teleoconch by slight varix, 1.1-1.2 whorls. First whorl 0.25-0.26 mm. in di- ameter. Penultimate and last whorls: flattening below suture and slight shoulder (PI. 56, figs. 1-3); commonly: slight spiral bulge around um- bilical area (Pl. 56, figs. 4, 9). Sculpture of sec- ond whorl: 3 spiral keels on first quadrant fol- lowing protoconch varix, the middle keel (on shoulder) the most prominent; keels less prom- inent on second to fourth quadrant, gradually be- coming spiral cords, with one or two more cords intercalated; very fine axial threads (PI. 56, fig. 5). On later whorls, ‘cords are reduced and broadened to interspaces between spiral sulci; 8- 12 sulci above suture on penultimate whorl. Last whorl of small shells: 25-28 spiral sulci between suture and base. Sulci absent at and near peri- phery of large shells, and wholly absent near outer lip where surface is smooth except for ax- ial growth lines; sulci fairly regularly to irregu- lary spaced (fairly uncommonly, in closely-spaced pairs), especially variable below and near suture; termination of sulci commonly abrupt. Surface shiny. Colors: pale pinkish brown or orange- brown, and white. Patterns: no demarcated sub- sutural area; on middle whorls: irregular axial bars at shoulder (0-10 per whorl), and near and [04-167] 58 Gabrielona above periphery (7-17 per whorl), developing into wavy bands; on last whorl large shells: ax- ially aligned, broadly wavy brownish bands aris- ing from suture and extending to base, 6-11 on last whorl, commonly coalesced into irregular spiral areas on shoulder and on base, and com- monly disjunct subperipherally; umbilical area: white, with 5-9 (usually 6) axial or very steeply descending brownish bands extending from mid- dle of base; fairly rarely: all brownish marks very pale and peripheral area uniformly whitish (PI. 54, fig. 4). Aperture slightly more rounded than in other Recent species (less constricted near su- ture); outer lip and callus on upper parietal area fairly thick to thick; palatal denticle on most large shells, commonly wide and faint; apertural ridge (ar) shallow in aperture; palatal sulcus near middle of aperture (lacking in smallest shell); upper columellar denticle (cd;) absent; lower columellar denticle (edz) present, commonly wide (Pl. 56, figs. 4, 9). Columellar lip not thickened, and steep slope into umbilical channel but no escarpment; escarpment to left of umbilical channel tairly prominent, arising fairly high to high off outer edge columellar lip; broad area to left of this escarpment, commonly with several parallel threads; umbilicus fairly wide to wide. Plate 55. Gabrielona sulcifera Robertson. Off northwestern Cuba. Abnormal shell. All x20. Robert Robertson Phasianellidae Abnormal shell (Pl. 55)—The single empty shell from 287 fathoms off northwestern Cuba (USNM no. 94974) differs strikingly from all those from Antigua, but seems to be conspecific. The proto- conch and beginning of the second whorl are normal, but the surface is increasingly corroded as far as a growth line in the first quadrant of the third whorl. Thereafter, the external surface is smooth and shiny and lacks sulci. Another growth line is at the beginning of the second quadrant of the third whorl. I conclude from the growth-lines, which are much more clear-cut than on any other Gabrielona observed, that growth of this shell was abnormal, perhaps be- cause the animal lived in an unusual habitat (depth?). Other differences from the Antiguan specimens: spire lower (spire angle ca. 115°); width exceeds length (resembles G. pisinna in outline); thick; coloration dark brick red and white; dark areas large, with banding complex and greatly coal- esced; apertural ridge (ar) fairly deep in aper- ture; slight trace upper columellar denticle (ed,); shallow slope from columellar lip into wide um- bilical channel; no trace escarpment to left. Shell measurements (mm. )— length width no. whorls 2.42 2.27 4.3 largest (Antigua) 2.13 2:20 3.8 abnormal (N.W. Cuba) 1.98 1.93 4.0 average; holotype 1.33 1.17 She) smallest (St. Croix) Operculum (Pl. 56, figs. 6-8)—The single oper- culum available is different from all other known Gabrielona opercula in having a prominent spiral cord at the inner edge of the marginal spiral ridge. The operculum of G. sulcifera is almost as thick as those of G. raunana, and the opercular whorl counts of these two species are also sim- ilar. In outline, the operculum of G. sulcifera is slightly more rounded than those of all the other Recent species—a consequence of the more rounded aperture. In the central region, a faint spiral ridge separates the smooth outer spiral area from the smooth central area. The callus is thin, revealing some of the spiral suture at the surface. The non-spiral columellar edge is steeply beveled. The spiral edge seems to be corroded at the external surface, and the sharp crest of the spiral ridge may therefore be an erosional fea- ture. Radula (Pl. 57; 1 studied)—Attains length of 0.7 mm., width of 0.13 mm., and with as many as 30 [04-168] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 Gabrielona — 59 fairly strongly curved transverse rows of teeth (including a few nascent rows). Central monocus- pid, with the small anterior portion of the base in a cleft between the bases of the innermost pair of laterals; the remainder of the base of the central overlapping these (where they are juxta- posed posteriorly). Five pairs of laterals, each of these (like the central) with a small distal por- tion that is monocuspid; the outer edge of each base partly overlaps the base lateral to it. As many as 34 pairs of marginals (lowest count 26), the innermost large and massive with lobed cusps, the distal portions gradually becoming smaller and more slender outwards, and with finer denticulations that finally are absent altogether on the outermost teeth. Synonymy— 1889 Phasianella (Eucosmia) brevis “Orbigny” Dall [Eucos- mia brevis in plate caption] (not P. brevis Orbigny, 1842). Bull Mus. Comp. Zool. 18: 30 [listed], 351, Plate 56. Gabrielona sulcifera Robertson. Figs. 1-3. Outlines of large shells, showing variation, all x17. Fig. 1. Abnormal shell from off northwestern Cuba. Fig. 3. Unusually high- spired shell. Fig. 4. Holotype, showing position of sulci, x33. Fig. 5. Apex, x60; p, protoconch. Figs. 6-8. Operculum of pl. 19, fig. 10b [not North Carolinian specimens, which are Tricolia thalassicola Robertson (deep water form) and perhaps other species of Tricolia}. 1918 Eucosmia brevis (“Orbigny”)Cossmann (not P. brevis Orbigny). Essais Paleoconch. Comp., Paris, livr. 11, p 162, fig. 55 [outline copied from Dall, 1889]. 1958 Gabrielona brevis (“Orbigny”) Robertson (not P. brevis Orbigny, 1842). Johnsonia 3. (37): 257-260, pl. 138, fig. 2 [radula], pl. 139, figs. 3-4 [operculum], pl. 140, figs. 2-3, pl. 141 [not pl. 142, fig. 1]. Types—The holotype (PI. 54, fig. 4), the live-col- lected specimen from Antigua, is at the United States National Museum (no. 500636), Washing- ton, D.C. Of the original 11 Antiguan paratypes, 7 are still at USNM (no. 659066), 2 are at the Museum of Comparative Zoology (no. 188356), 1 is at the Academy of Natural Sciences of Phila- delphia (no. 302625), and 1 (PI. 54, fig. 3) was lost. The smallest paratype, fromm St. Croix, is in Mr. Usticke’s private collection. The ab- normal shell from Cuba is not a paratype. Derivation of new name—Latin, sulcifer, bearing furrows. holotype. Internal surface, longitudinal section, and external surface (respectively). All x33; c, lower edge of callus. Fig. 9. Aperture, x33; ar, apertural ridge; ecd,, lower (inner) colu- mellar denticle; pd, palatal denticle; ps, palatal sulcus; uc, umbilical channel. [04-169] 60 Gabrielona Nomenclature—Following Dall (1889), who first applied the name Phasianella brevis Orbigny, 1842, to the abnormal Cuban shell discussed and retigured here (Pl. 55), I misapplied this name to this species and transferred it to Gabrielona. Subsequent study of the holotype (figured speci- men) of P. brevis at the British Museum (Nat. Hist.), no. 1854.10.4.282, has shown that this is a depauperate Tricolia. This holotype, inadequately illustrated in Robertson (1958, pl. 142, fig. 1), will be treated in detail and refigured elsewhere. Robert Robertson Phasianellidae Locality records (see map, Pl. 53, circular black spots)— CUBA: off Bahia Honda, Pinar del Rio (23°2’N.; 83°13’W.; 287 fms.; Blake Sta. 21 [1877-78]; USNM). VIRGIN ISLANDS: Christiansted Harbor, St. Croix (dredged 15 ft [1 dead]; G. N. Usticke). LESSER ANTILLES: English Harbour, Antigua (1918, J. B. Henderson, Jr., USNM, MCZ, ANSP). Erroneous locality record—Arenas de la Chor- rera, Habana, Cuba (Robertson, 1958, pp. 259- 260, as G. brevis) [a juvenile Tricolia]. als. The cusps of the central and laterals are stippled. From the holotype, Antigua, West Indies. x1500. Modified from Robertson (1958, pl. 138, fig. 2, as Gabrielona brevis). Plate 57. Gabrielona sulcifera Robertson. Half of one trans- verse row of radular teeth, showing (from left to right) the central between the innermost pair of laterals, the remaining four laterals on the right side, and the entire row of margin- [04-170] May 30, 1973 EXCLUDED SPECIES “Gabrielona” bruscasensis Weisbord, 1962 (PI. 58) Range—Upper Miocene or Pliocene (or younger?): known only from one locality in the Playa Grande Formation (Maiquetia Member), northern Venezuela. Remarks—This species was described from a single poorly preserved shell with a badly broken columellar area. Even generic identification has been a problem but the shell does seem to be a phasianellid. However, it cannot be a Gabrielona because the slight axial ridge revealing the posi- tion of the withdrawn operculum is detectable, and this is fairly deep in the aperture. I detected no palatal sulcus or apertural denticles, and the part remaining of the umbilical area indicates that there was no Gabrielona-like umbilical channel. The outer lip is more prosocline than in any true Gabrielona. I conclude that G. brusca- sensis probably is a young Tricolia. The specimen lacks most of the characters dis- tinguishing species of Tricolia. The apex is in such poor condition that the whorls cannot be counted accurately and the first whorl cannot be measured. The shell is suffused with pale brown- ish pink, but no color pattern is detectable. Sim- ilar-sized shells of T. affinis cruenta Robertson differ in outline (are more elongate, with a more obtuse apex) and lack the umbilicus. G. brusca- sensis is here considered a nomen dubium. Synonymy— 1962 Gabrielona bruscasensis Weisbord, Bulls. American Paleo., 42(193): 111, pl. 8, figs. 5-7 (Quebrada las Bruscas [Playa Grande Formation], Distrito Feder- al, Venezuela). Type —The holotype is at the Paleontological Research Institution (no. 26056), Ithaca, New York. Plate 58. “Gabrielona” bruscasensis Weisbord, probably a young Tricolia. Upper Miocene or Pliocene [?], northern Venezuela. Holotype. Both x20. INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 Gabrielona 61 “Gabrielona” sphaera Weisbord, 1962 (Pl. 59) Range—Pliocene (or younger?): known only from three nearby localities in the Mare Forma- tion, northern Venezuela. Remarks—This is a rissoacean. Although re- sembling in outline a high-spired Gabrielona, it cannot be one because the shell (for its size) is thin, the periphery of the last whorl is lower than in any known Gabrielona, the aperture is elon- gate-ovate, there are no apertural denticles, and neither is there a palatal sulcus. There is a wide umbilicus and a broad umbilical channel that is slightly angled at its lower left margin. The holo- type has 3.4 whorls and the first whorl is 0.23 mm. in diameter. The callus on the upper parie- tal area is incomplete medially. In coloration, the shell is pale (faded?) amber and whitish near the umbilicus and on the last part of the last whorl where there are irregular amber markings. At the apex, the beginning of the suture is tinged with dark amber. Synonymy— 1962 Gabrielona sphaera Weisbord, Bulls. American Paleo., 42(193): 109-111, pl. 8, figs. 1-4 (near Quebrada Mare Abajo [Mare Formation], Distrito Federal, Venezuela). Types—The holotype is at the Paleontological Research Institution (no. 26054), Ithaca, New York. So also is the single paratype (no. 26055) distinguished by Weisbord among eleven other specimens identified with G. sphaera. Plate 59. “Gabrielona” sphaera Weisbord, a rissoacean. Plio- cene [?], northern Venezuela. Holotype. Both x20. [04-177] Published by THE DELAWARE MUSEUM OF NATURAL HISTORY Box 3937, Greenville, Delaware 19807, U.S.A. [04-178] INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 June 1, 1973 {replacing vol. 2, no. 12, pp. 509-510; see stars] Tectariinae 63 trations of these morphological differences in Rosewater, 1970, and in this paper). Most Tectarius s.s.and Echininus s.s. are living today only in the East Indian area. With the exception of Tectarius grandinatus whose range extends eastward to Polynesia, all other species are inhabitants of the raised, weathered coral reef shorelines found in the Western Pacific Arc (per- sonal observations, 1970). It can only be assumed that this niche provides the requirements essen- tial for the existence of these species:as they are to be found nowhere else. The fossil record provides very few clues to the origin of these groups. There are only three Terti- ary fossil species described from the Indo-Pacific and these probably belong in three separate ge- neric taxa. The oldest of these, T. songoense Martin, from the Upper Eocene of Java, probably represents nearly the earliest appearance of Tec- tarius. AS mentioned in Part I, littorinid fossils are exceedingly difficult to separate from Trochi- dae and Turbinidae, and this is no less true for Tectarius and Echininus. Probably, however, these groups made their appearance in the early Tertiary within the region where they have devel- oped, and with the exceptions of Echininus (Tec- tininus) nodulosus and Tectarius (Cenchritis) muricatus, both of the western Atlantic, they have remained there. Opercula The opercula of Indo-Pacific Littorinidae re- quire special comment (see pl. 389). All are made up of conchiolin, and those of members of the subfamily Littorininae, including Littorina, its subgenera, and Nodilittorina are usually pauci- spiral and rather oval in shape with the nucleus at the side and nearer one end (oligogyrous spiral tvpe of Fretter, et al., 1962, pp. 79,80). In the Echinininae, the basic plan of the operculum differs from that of other littorines. It is the type D Plate 389. Opercula of Littorinidae and Trochidae. Fig. A. Paucispiral operculum of Littorina (Littorinopsis) scabra (Linne) from Mokuoloe Id., Kaneohe Bay, Oahu (USNM 339388). Fig. B. Mesospiral operculum of Tectarius rusticus (Philip- pi) from Troughton Chain, northern Western Australia (WAM 1787-69); Fig. C. Multispiral operculum of Echininus cumingi (Phi- lippi) from near Davao City, Mindanao, Philippines (WAM 1566-70); Fig. D. Multispiral operculum of Trochus niloticus Linne, from Makuluva, Viti Levu, Fiji (USNM 531827). Lines under each figure represent 5 mm.; stippled areas are thickened and dark-brown in color; non-stippled areas are light horn color and transparent. of operculum which is called in other groups, such as Trochidae, a miultispiral operculum (polygyrous spiral type of Fretter, et al., ibid.) although not so extreme as that figured by Fretter (ibid., p. 80, fig. 43A; also see our pl. 389, fig. D.). The operculum is circular in outline and moder- ate to small in size. The nucleus is decidedly central in location and growth proceeds outward from the center in multiple, fairly evenly spaced gyrations (pl. 389, fig. D). The operculum in Tec- tarininae (fig. B) is intermediate in form between Explanation to plate 388 (opposite page) Figs. 1,2. Tectarius grandinatus (Gmelin) from Palmerston Atoll, Cook Islands (USNM 685165) Figs. 3,4. Tectarius tectumpersicum (Linné). Fig. 3, from Stirling Isle, Treasury Ids., Solomon Islands (USNM 600370); Fig. 4, from “East Indies” (USNM 131450). Figs. 5-7. Tectarius pagodus (Linne). Fig. 5, from “East Indies” (USNM_ 18966); Fig. 6, from the Philippines (USNM); Fig. 7, a young specimen from Polillo, Philip- pines (USNM 311141). Figs. 8.9. Tectarius rusticus (Philippi) from Buccaneer Archipelago, Western Australia (USNM 684713). Figs. 10,11 Tectarius coronatus Val. Fig. 10, from Pacific (USNM 304587); Fig. 11, from Davao Bay, Mindanao, Philippines (USNM 654034). Figs. 12,13. Echininus cumingi cumingi (Philippi) from Hervey Ids., Cook Ids. (USNM 42452). Figs. 14,15. Echininus cumingi spinulosus (Philippi) from Kadena Circle, Okinawa, Ryukyu Ids. (USNM 664658). Figs. 16,17. Tectarius (Cenchritis) muricatus (Linné), from Boca de Camarioca, Matanzas, Cuba (USNM 599944). Figs. 18-20. Echininus (Tectininus) nodulosus (Pfeiffer). Fig. 18, from Hog Island, Bahamas (USNM_ 603911); Figs. 19,20, from Mujeres Harbor, Quintana Roo, Mexico (USNM 662308). (05-443a] 64 Joseph Rosewater Littorininae and Echinininae, being large and rather rounded in outline, with a slightly acentric nucleus and having a number of gyrations more than the paucispiral type but less in number than the multispiral type. The Tectariine opercular type is here termed the mesospiral or mesogyrous spiral tvpe. The presence of the three opercular types in Littorinidae may be considered to have evolutionary significance, and possibly is related to selection for a better aperture sealing mecha- nism in animals which have considerable vertical distribution on the shore line: in order, proceed- ing from low toward higher shore habitats— Littorininae, Tectariinae, Echinininae. Reproduction To my knowledge nothing is known concern- ing reproduction in either Tectariinae or Echin- ininae with the exception of Tectarius (Cen- chritis) muricatus (Linné) which produces a pe- lagic capsule (see Lebour, 1945, and Lewis, 1960, references in Rosewater, 1970 p. 05-276). Field and laboratory studies are needed to discover details of the life histories of the remaining spe- cies. However it is likely that most of these snails also produce eggs encased in pelagic capsules which undergo development in the sea. Abbott (1954) noted that Lebour (ibid.) stated that some of the Bermuda littorinids that live above high tide line migrate to the water to spawn. It is suspected that this also is the case with many of the Indo-Pacific species. Acknowledgments The persons and institutions acknowledged in Part I of this study (see Rosewater, Indo-Pacific Mollusca, vol. 2, no. 11, p. 425) also are thanked here. In addition, I acknowledge the following for their help in making possible the examination in the field of most of the species of Tectarius and Echininus during the National Geographic Socie- tv—Mariel King Memorial Expedition to the Mo- luceas Islands, Indonesia, May to July 1970: the late Mariel King, Mrs. Grace King, T. H. Richert, C. Beal, C. M. Burgess, B. R. Wilson, and the National Geographic Societv. The Government of Indonesia graciously provided clearance for the vessel Pele to work in the Moluccas Islands. Mr. Kasim Moosa and Mr. Sukarno, both of the Insti- tute for Marine Research, Djakarta, accompanied the expedition and provided assistance of many kinds. Littorinidae List of Recognized Taxa Below is a list of the Tertiary fossil and Recent species herein recognized as belonging in the subfamilies Tectariinae and Echinininae. The few fossil taxa are preceded by a dagger [#]. Family Littorinidae Gray, 1840 Subfamily Tectariinae, new subfamily GENUS Tectarius Valenciennes, [1832] Subgenus Tectarius Valenciennes, [1832] coronatus Val., 1832. Type. Recent, wes- tern Pacific grandinatus (Gmelin, 1791). Recent, Pacific islands pagodus (Linné, 1758). Recent, western Pa- cific tectumpersicum (Linné, 1758). Recent, wes- tern Pacific rusticus (Philippi, 1846). Recent, northern Australia t songoense (K. Martin, 1931). Eocene, Java. Subgenus +Subditotectarius Ladd, 1966 trehderi Ladd, 1966. Type. Miocene, Mar- shall Islands. Subgenus Cenchritis von Martens, 1900 muricatus (Linne, 1758). Type. Recent, tropical western Atlantic. Subfamily Echinininae, new subfamily GENUS Echininus Clench and Abbott, 1942 Subgenus Echininus Clench and Abbott, 1942 cumingi cumingi (Philippi, 1846). Type. Re- cent, western Pacific cumingi spinulosus (Philippi, 1847). Recent, western Pacific + adelaidensis (Cotton, 1947). Pliocene, South Australia. Subgenus Tectininus Clench and Abbott, 1942 nodulosus (Pfeiffer, 1839). Type. Recent, tropical western Atlantic. [05-44-4a] INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 June 1, 1973 [replacing vol. 2, no. 12, pp. 511-512; see stars] Tectariinae 65 Selected Bibliography Abbott, R. T. 1954. Review of the Atlantic Periwinkles, Nodilittorina, Echininus, and Tectarius. Proceedings of the United States National Museum, vol. 103, no. 3328, pp. 449-464. Argenville, A. J. D. d’. 1742. L’histoire naturelle —La Litho- logie et la Conchyliologie —par—M. [A. J. D. d’ Argenville] de la Société Royale des Sciences de Montpellier. Paris. Bruguiere, M. 1792. Encyclopédie Méthodique, Paris, vol. 1, p. 930. Clench, W. J. and R. T. Abbott. 1942. The Genera Tectarius and Echininus in the Western Atlantic. Johnsonia, vol. 1, no. 4, pp. 1-4. Dance, S. P. 1967. Report on the Linnaean Shell Collection. Proceedings of the Linnean Society of London, vol. 178, no. 1, pp. 1-24, 10 pls. Dell, R. K. 1964. Marine Mollusca from Macquarie and Heard Islands. Records of the Dominion Museum, vol. 4, no. 20, pp. 267-301. Deshayes, G. P. 1830. Encyclopédie Meéethodique, Paris, vol. 2, p. 184. Dodge, H. 1959. A Historical Review of the Mollusks. of Linnaeus. Bulletin of the American Museum of Natural History, vol. 118, Article 5, pp. 211-257. Fretter, V., and A. Graham. 1962. British Prosobranch Mol- luscs. Ray Society, London, xvi + 755 pp., 317 figs. Habe, T. 1951. Littorinidae in Japan (1). Hlustrated Catalogue of Japanese Shells, no. 14, pp. 87-93. Habe, T. 1961. Coloured Ilustrations of the Shells of Japan (II). Hoikusha Publishing Co., Ltd., Osaka, 183 pp., 66 pls. Habe, T. 1964. Shells of the Western Pacific in Color, Vol. I. Hoikusha Publishing Co., Ltd., Osaka. 233 pp., 66 pls Kaicher, S. D. 1956. Indo-Pacific Sea Shells. Section 3. Lit- torinacea, etc. Privately Printed, Washington, D.C., 8 pls. and captions. Keen, A. Myra. 1966. Tectarius (Mollusca: Gastropoda): Re- quest for Validation in its Accustomed Sense. Z.N. (S.) 1754. Bulletin of Zoological Nomenclature, vol. 23, part 4, pp. 179-180. Kesteven, H. L. 1903. Notes on Prosobranchiata, No. II. Littorinacea. Proceedings of the Linnean Society of New South Wales, 1902, part 4, pp. 620-636. Kira, T. 1959. Coloured Illustrations of the Shells of Japan. Revised Edition, Hoikusha Publishing Co., Ltd., Osaka, ix + 239 pp. Kira, T. 1962. Shells of the Western Pacific in Color. Hoikusha Publishing Co., Ltd., Osaka, 224 pp. Klein, J. T. 1753. Tentamen Methodi Ostracologicae sive Dispositio Naturalis Cochlidum et Concharum, p. 25. Lug- duni Batavorum. Melville, R. V. and W. E. China. 1969. Opinion 871. Tectarius Valenciennes, [1832] (Gastropoda): Validated Under the Plenary Powers. The Bulletin of Zoological Nomenclature, vol. 25, part 6, pp. 214-215. Morch, O. A. L. 1852. Catalogus Conchyliorum Quae Reliquit D. Alphonso D’Aguirra et Gadea Comes de Yoldi; fascicle 1, p. 45. Powell, A. W. B. 1951. Antarctic and Subantarctic Mollusca: Pelecypoda and Gastropoda. Discovery Reports, vol. 26, pp. 47-196, text figs., 6 pls. Prashad, B. 1925. Respiration of Gastropod Mollusks. Pro- ceedings of the Twelfth Pacific Science Congress, pp. 126-143. Rosewater, J. 1970. The Family Littorinidae in the Indo- Pacific. Part I. The Subfamily Littorininae. Indo-Pacific Mollusca, vol. 2, no. 11, pp. 417-506, 64 pls. Rumphius, G. E. 1705. D’Amboinsche Rariteitkamer. Am- sterdam, 340 pp., 60 pls. Sherborn, C. D. and B. B. Woodward. 1901. Bibliographical Notes. XXVII. The Dates of Humboldt and Bonpland’s “Voyage”. Journal of Botany, June, pp. 1-4. Troschel, F. H. 1856-1863. Das Gebiss der Schnecken, vol. 1, Berlin, pp. vii + 252, pl. 1-20. Watson, R. B. 1886. Report on the Scientific Results of the H.M.S. Challenger, vol. 15, part 42, p. 576. Wenz, W. 1938. Handbuch der Paliozoologie, vol. 6, part 1, lfg. 3, pp. 241-480, figs. 472-1235. Wimmer, August. L880. Sitzungsberichte der Mathematisch- naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften, Wien, I. Abth. vol. 80, pp. 496, 514. [05-447a ] 66 Joseph Rosewater Littorinidae Key to the Tectariinae and Echinininae The following key is to the genera and sub- only, since Western Atlantic taxa are not treated in genera of these two subfamilies. It is based upon detail in the present paper. For a key to the the shell and externally observable characters. Littorininae, see vol. 2, no. 11, p. 430 [p. 05-278]. Page numbers are given for Indo-Pacific groups LA OMe Wi DINER. 0 4 ox oot ered. eed 4ae 6S ERODES ee RRA ae aes 2 Lb: Sivell wet sa Bia ccs cs Gott ae ba eee eh We ee Rd Se Ge dew ed Gels ha ee dma ed 3 2a Shell spinose, with partially open spines, shell about as wide as high, operculum pipe iS (see Pi SSO). oe es eka eo bea eh Oa eeds Homa dad Echininus p. 526 2b Shell not spinose, nodulose, higher than wide, operculum not multispiral . .. Cenchritis 3a Shell spinose, operculum mesospiral. Tectarius .......0.00 0000 ce ee ees p.o13 Sb Shell moculose,70S6il i oo ioc oo kaka he Pe eee dR KK RS Subditotectarius p. 524 3c Shell moderately spinose, operculum multispiral................005. Tectininus ie Plate 390. Type-species of Genera and Subgenera of Tectari- Miocene, Marshall Islands (Holotype, USNM 648342; 2.8 inae (Figs. 1-9) and Echinininae (Figs. 10-15) illustrating x 2.4 mm.). sculpture, arrangement of spines, and presence or absence of Figs. 7-9. Tectarius (Cenchritis) muricatus (Linné); umbilici. Matanzas, Cuba (USNM 599944, 26.1 x 17.9 mm.). Figs. 10-12. Echininus (Echininus) cumingi (Philippi); * Figs. 1-3. Tectarius (Tectarius) coronatus Val.; Davao Bay, Cook Islands (USNM 42452a; 17.2 x 17.5 mm.). Mindanao, Philippines (USNM 654034; 23.1 x 17.8mm.). Figs. 13-15. Echininus (Tectininus) nodulosus (Pfeiffer); Figs. 4-6. Tectarius (Subditotectarius) rehderi Ladd; early Cozumel Id., Mexico (USNM 662806; 14.4 x 12.2 mm.). [05-448a ] * INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 June 1, 1973 [replacing vol. 2, no. 12, pp. 513-514; see stars] Tectarius 67 Subfamily Tectariinae, new subfamily Genus Tectarius Valenciennes, [1832] Type: Tectarius coronatus Val., 1832 The genus Tectarius sensu lato includes the nominate subgenus, Tectarius, whose type- species T. coronatus Val. (- T. papillosus ‘La- marck’ of authors) has one of the more conserva- tively sculptured shells of the group. Tectarius sensu stricto is a wholly Indo-Pacific group, hav- ing its present population center in the Western Pacific Arc. The monotypic subgenus Cenchritis von Martens contains only TJ. (C.) muricatus (Linné), of the tropical western Atlantic (see pl. 388, figs. 16, 17). Subditotectarius Ladd, 1966, is monotypic for the fossil T. (S.) rehderi Ladd, of the Miocene of the Marshall Islands. Only spe- cies belonging to Tectarius s.s. and Subditotec- tarius will be considered here. Tectarius appears more closely related to Lit- torina than to Echininus because of greater simi- larities in morphology, general shell characters, the absence of a truly multispiral operculum, a broader, less modified central radula tooth, and the usual lack of an openly umbilicate shell. Subgenus Tectarius sensu stricto Moderately large, pyramidal to turbinate, non-umbilicate littorinids with from rather strongly spinose to nodulose or papillose shells; generally living at or above high tide line. Radula littorinoid, the central tooth somewhat narrowed, the lateral tooth with an embayment and usually developing a medial vertical ridge or partition. In males the penis is large and well-supplied with glands along most of its lateral edge, the remain- der papillose, and with an open but deeply folded seminal duct. Operculum rounded, mesospiral (see Opercula in Introduction). Aperture plicate within; with a columellar swelling or tooth. Synonymy— 1798 Cidaris Réding, Museum Boltenianum, part 2, p. 84; type-species by subsequent designation, Herrmann- sen, 1847: Trochus pagodus Linné; non Cidaris Leske, 1778, nor Swainson, 1840. [1832] Tectarius Valenciennes, Coquilles, in Humboldt and Bonpland. Voyage aux régions équinoxiales du Noveau Continent, Observations de Zoologie, vol. 2, p. 271; type-species by subsequent designation Clench and Abbott, 1942: Trochus coronatus Valenci- ennes. 1839 Pagodus Gray, in Molluscous Animals: The Zoology of Captain Beechey’s Voyage, p. 141; type-species by Monotypy and by absolute tautonymy, Monodonta pagodus Lamarck [= Tectarius pagodus (Linné]. 1840 Pagodella Swainson, A Treatise on Malacology, pp. 207, 219, 221 [refers to Pagodella echinata, nomen nudum] 351; refers to P. major Martini - Chemnitz, pl. 163, figs 1541, 1542 [= Tectarius pagodus (Linné)] and to T. tectumpersicum ibid., fig. 1543, 1544; type-species here designated: Tectarius pagodus (Linneé). 1840 Echinella Swainson, ibid., pp. 207, 221, 352; refers to E. granulata Swainson [nomen nudum] and to E. coro- naria, Tableau Encyclopédique et Methodique, pl. 447, fig 6 [=Mondonta coronaria Lamarck = Tectar- ius grandinatus (Gimelin)]; type-species by monotypy, Tectarius grandinatus (Gmelin) [also see Clench and Abbott, 1942]; not Echinella Bory St. Vincent, 1824. 1846 Fectaria Philippi, Abbildungen und Beschreibungen Conchylien, Vol. 2, Litorina, p. 139; used in combina- tion Fectaria pagodus, error for Tectarius Valencien- nes: 1858 Hamus ‘Klein’ H. & A. Adams, The Genera of Recent Mollusea, vol. 2, p. 656, refers to H. and A. Adams, 1854, vol. 1, p. 315; type-species here designated, Hamus pagodus (Linne) [=Tectarius pagodus (Linné)]; not Hamus ‘Klein’ R. B. Watson, 1886 {=Trochidae]. 1899 Echinellopsis Rovereto, Atti della Societa Ligustica di Scienze naturali e geografiche, vol. 10, p. 109; new name for Echinella Swainson, 1840, not Bory St. Vincent, 1524. Nomenclature—Due to similarities between the shells of Tectarius and some of the Trochidae, there has been a tendency for some of the former to be classified with the latter. This problem was discussed by Keen (1966) who recommended that the Zoological Nomenclature validate Tectarius with the type- Tectarius coronatus Valenciennes, [1832], i.e., in its accustomed sense. Her petition was granted in I.C.Z.N. Opinion 871 (Melville and China, 1969). It was assumed in this Opinion that the type-species of Tectarius, T. coronatus International Commission. on species Valenciennes, is a synonym of T. grandinatus * Gmelin. However, these are in actuality distinct and geographically isolated species. Another name which sometimes has been as- sociated with Tectarius that has an exceedingly long and complicated history is the genus Ha- mus. It was mentioned originally by Klein (1753) where its use was of course pre-linnaean. Bru- guiéere (1792) gave a brief description, referring to Klein, but listed no species. Deshaves (1830) [05-449] 68 Joseph Rosewater Littorinidae declared it “a forgotten genus’, indicating that he considered it unrecognizable. Mérch (1852)listed it, this time in the synonymy of Littorina Férus- sac, an invalid introduction (I.C.Z.N., Art. 11(d)). The first valid use of Hamus was not until H. & A. Adams (1858) used it as a senior synonym for Tectarius (see synonymy). It was later used by Wimmer (1880) and Watson (1886), the last being a taxon of Trochidae. I have designated as type- species of Hamus H. and A. Adams, 1858, H. pagodus (Linné) and consider this genus to be an absolute synonym of Tectarius Valenciennes. The use of square brackets surrounding the date for Tectarius Valenciennes, [1832] is reecom- mended by the International Code of Zoological Nomenclature in cases where the date of publica- tion of a name has been determined on the basis of external evidence (I.C.Z.N. Recommendation 92.A(3); also see Sherborn and Woodward, 1901; Keen, 1966; and Opinion 871). Tectarius coronatus Valenciennes, 1832 (PI. 388, figs. 10, 11) Range—Philippines and Indonesia. Remarks—The shells of well prepared and cleaned specimens of Tectarius coronatus tendto colorful for with the orange-pink coloration of the last two whorls contrasting with a purplish brown. subsutural band. These colors do not show well in all speci- mens, however, and are not very visible in un- cleaned The umbilicate common species of the southwest Pa- be quite Littorinidae, specimens. three large, non- cific may be distinguished by the number of major spiral rows of spines on the last whorl: 2 in pagodus; 3 in tectumpersicum; and 4 in coronatus. The closely-spaced stubby spines of coronatus also separate it from the other two. These characteris- tics do not of course help to distinguish it from T. grandinatus, but other characters and the Polyne- sian endemicity of the latter are helpful in this case (see Remarks under grandinatus). Habitat—Shore rocks and limestone cliffs 1-2 meters above high tide line (personal observa- tions, Davao, Philippines, 1970). Description—Shell reaching 39.7 mm (about I'/2 inches) in length, broadly conical in shape, average obesity about .76 (51 specimens range from .68-.82); heavily constructed, imperforate, and sculptured mature specimens moderately on most postnuclear whorls with four, fairly closely-spaced rows of stubby, rounded, often slightly upturned spines. External color generally yellowish white on early whorls, becoming pink- ish orange on penultimate and body whorls; area of most posterior (subsutural) row of spines usual- ly a contrasting purplish brown, and the same dark color may appear in lines and dashes inside outer lip of aperture; aperture tinted lighter pink- ish orange. Base flattened, sculptured spirally with nodulose cords, a larger separate row just below periphery of body whorl. Whorls 6-8, flat- sided excepting spines. Length of spire usually greater than half the length of shell. Spire convex, produced at an angle of from about 60-67°. Aper- ture rounded-squarish; outer lip thickly produced in mature individuals, strongly plicate within; plicae not reaching edge of aperture; outer lip tapering to a thin, crenulate edge; inner lip smooth posteriorly, often stained a deeper orange than rest of aperture, forming a tooth-like bulge anteriorly, near junction with outer lip near base of columella. Suture obscured by anteriormost row of spines of preceding whorl. Primary sculp- iural feature is the four spiral rows of spines. Spines not particularly aligned axially, although anteriormost 2 rows more so than others; from 17-23 spines per row on body whorl, bases of anteriormost 3 rows of spines joined by low spiral carinae. Posteriormost-but-one (3rd) row of Plate 391. Tectarius coronatus Figs. 1,2. Turbo rugosus Wood, lectotype, BM(NH) 1968370, 28.4 * 22.3 mm. Figs. 3,4. Tectarius coronatus Valenciennes, Holotype, MHNP (“Acapulco” [Luzon, Philippines]) 32.9 x 25.7 mam. [05-450a] * INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 June 1, 1973 [replacing vol. 2, no. 12, pp. 515-516; see stars] Tectarius 69 spines protrudes farthest on spire whorls, but this distinction largely lost on more mature whorls; posteriormost (4th) row of spines obscured by subsutucal purplish brown color band especially on penultimate and body whorls. Secondary spiral sculpture, between each row of spines, con- sisting of raised cords, and overall spiral sculp- ture of microscopic threads. Axial sculpture con- sists of irregular flaky lines of growth. Operculum moderate in size, circular, an average One measur- ing about 7 mm. in diameter, thin, light-brown with a dark-brown center, paucispiral, nucleus about central. Periostracum not evident. Nuclear whorls at least partially decollate in all specimens examined, about 2, smooth, grayish white, first postnuclear whorls showing early signs of spiral striae and becoming nodulose. Radula littorinoid, 2-1-1-1-2; lateral tooth with a vertical partition and an embayment typical of Littorinidae. Ani- mal moderately large, littorinoid; penis large, muscular and apparently highly extensible; semi- nal groove in deep fold running along medial edge to tip; tip vermiform, covered with papillae; lateral edge of penis supplied with large number of glands not extending onto vermiform tip. Re- productive data and life history unknown. Measurements (mm) (all Philippines)— length width No. locality whorls 39.7 27.6 7+ Cadao Id., Naro Bay, Masbate 35.4 25.2. 7+ Cadao Id., Naro Bay, Masbate 30.1 22.4 7+ San Miguel Bay, Ticao 95.9 D2, 8 Batag Id., Samar 23.9 16.3 7+ Bongao Channal, SW Sanga Sanga Id., Sulu Archipelago 21.0 15.3 7+ Borongan, E side Samar L725 130 7+ Papahag Id., Tawi Tawi Group 15.0 12.0 6+ 13.5 10.8 7+ 12.6 9.8 6+ Lee. 5.4 6+ 5.4 6.8 5+ Borongan Village, E side Samar Papahag Id., Tawi Tawi Group Papahag Id., Tawi Tawi Papahag Id., Tawi Tawi Papahag Id., Tawi Tawi Synonymy— ——— Monodonta papillosa of authors, not M. papillosa Lamarck, 1822 [=Tectarius tectumpersicum (Linné, 1758) ]. 1828 Trochus rugosus Wood, Supplement to the Index Tes- taceologicus or a Catalogue of Shells, British and Foreign, pl. 5, Trochus, fig. 7 (no locality given; Mindanao, Philippines, here selected); lectotype in BM(NH) 1968370, length 28.4 mm, width ( ca. ) 22.3; not Litorina rugosa Menke, 1843 [= Nodilittorina aus- tralis (Gray, 1926)]. Not Roding, 1798; Brown, 1815. 1832 Tectarius coronatus Valenciennes in Humboldt and Bonpland, Voyage aux régions equinoxiales du Noveau Continent, vol. 2, Coquilles, p. 271 (Acapulco lin error] locality here corrected to Luzon, Philip- pines); Holotype in MHNP. 1846 Litorina papillosa elegans Philippi, Abbildungen und Beschreibungen Conchylien, vol. 2, p. 140, Litorina, pl. 2, figs. 5, 7 (precise locality not given); figured specimens from Cuming Collection BM(NH) [not seen during 1968 visit] and Saul Collection, Cam- bridge Museum. 1846 Litorina papillosa quadriseriata Philippi, ibid., p. 140, Litorina, pl. 2, fig. 2 (Zanzibar [in error] locality here corrected to Luzon, Philippines); type-specimen may be in BM(NH) [not seen during 1968 visit]; refers to “Trochus rugosus Wood Suppl. t.5, f. 7”. Types—Although the name T. papillosus has been applied to this species (see Kaicher, 1956) the type-specimen of that species in the Geneva Museum is unquestionably T. tectumpersicum Linné, and papillosus is, therefore an absolute synonym of tectumpersicum (q.v.). It also has been referred to as Echinellopsis grandinatus (Habe, 1961, p. 20; 1964, p. 28, both pl. 9, fig. 30) which is an error of nomenclature for T. rugosus. The first available name is Tectarius coronatus Val. [1832], the holotype of which is in the Paris Museum. The figured specimens of Philip- pis elegans and quadriseriata may be in the BM (NH) and/or the Cambridge Museum. They were not discovered by me at the BMand may be lost. Philippi’s figures are quite adequate for the interpretation of the species and may be con- sidered as representative of the lectotypes: ele- gans, pl. 2, fig. 7; quadriseriata, pl. 2, fig. 2. \Oo0° 120 140° 16¢ T T BONIN Ss Jima » Morcus ow \ = * MARIANA x * ANDAMAN anaes | iE ’ BHILIPPINE £ Guns iS . Eniwetok Yop ith; ] NICOBAR ; i * 08 blau 4 F : id res ZEYLON CAROLINE iss €OCO6 - KEELING ATOLLS ¥ | 1 | 1 TOo° 120° 140° 160 Plate 392. Geographic distribution of Tectarius coronatus x (Wood) in the Philippines and Indonesia. [05-45 1a] 70 Joseph Rosewater Records—PHILIPPINES: Port oe Mindoro; San Miguel Bay, Ticao Id.; Cadao Id., Naro Bay, Masbate (all USNM); Borongan Village ( (USNM, Del. Mus. ee ANSP, MCZ): Babee ha both Samar (USNM); Samal Id., Davao Bay (MCZ. USNM, WAM): Zamboanga, both Mindanao (ANSP, Del. Mus. N.H.); Jolo Id. (MCZ); Tabawan Id. (ANSP); Papahag Id. (USNM); Bongao Channel, SW Sanga Sanga Id., all Sulu Archipelago (ANSP). INDONESIA: Buka Buka Id.. Gulf of Tomini, Celebes (USNM). Tectarius grandinatus (Gmelin, 1791) (Pl. 388, figs. 1,2; pls. 393-395) Range—The Cook, Society, Tuamotu and Gambier Islands, southeastern Polynesia. Remarks—Tectarius grandinatus, an inhabitant of southeastern Polynesia, apparently is ge- ographically isolated from the several other mem- bers of the genus Tectarius in the Indo-Pacific, the others being found in the East Indies. It is quite distinct, morphologically, from pagodus and tectumpersicum, but, interestingly, is very ¥& close in appearance to T. coronatus in general matters of sculpture and external anatomy of the animal. Superficially, grandinatus might be thought more closely related to Cenchritis muri- catus (L.) of the western Atlantic. However, the radula, penial anatomy (Abbott, 1954) and a close examination of shell sculpture and_ structure causes me to reject that theory: grandinatus has not been observed to be umbilicate, while muri- catus sometimes is; the aperture of muricatus is never thickened and plicate as it is in grandinat- us; the operculum of grandinatus is rounded with the nucleus near the center, that of muricatus is more oval with the nucleus near the side. The many similarities between coronatus and grandinatus together with their spatial isolation suggest the possibility that they may have evolved from the same species stock. Although a subspecific relationship may be indicated, the two are here considered to have developed full specific differentiation. Habitat—Usually occurs on low islands within the geographic range; on coral reef flats near the high tide line, on jagged pieces of raised reef. Description—Shell reaching 37.9 mm (about 1 1/9 inches) in length, elongate conical in shape, average obesity about .72 (44 specimens range from .62-.82); rather solidly and thickly con- structed for its size, imperforate; mature speci- mens with very deeply impressed suture and rounded whorls; sculptured with four spiral rows per whorl of stubby spines. External shell color yellowish white; often coated with what appears Littorinidae to be a light-yellow to brown or rather dark grayish brown periostracum which may wear thin especially at tips of spines; no definite color pattern apparent; occasional lines or splotches of medium brown present; upper (most posterior) part of aperture usually covered with a brown glaze. Base moderately flattened, sculptured spi- rally with nodulose cords. Whorls 7-9, moderately rounded. Length of spire usually considerably greater than half the length of shell. Spire convex, produced at an angle of from 58-60°. Aperture rounded; outer lip considerably thickened, pli- cate within, tapering to a thinner, crenulate edge. Inner lip with a thin, brown plage posteriorly, forming a tooth-like bulge anteriorly, near junc- tion with outer lip near base of columella. Suture often deeply impressed, typically forming a chan- nel between whorls. Primary sculptural feature is the four spiral rows of spines. Spines not regu- larly aligned axially, although anteriormost two rows more-so than others; from about 17-25 spines per row on body whorl; bases of anteri- ormost 3 rows of spines joined by low spiral carinae; bases of posteriormost row of spines usually separate; second from anterior row usual- ly the smallest. Secondary spiral sculpture be- Plate 393. Tectarius grandinatus (Gmelin, 1791). Figs. 1,2. Trochus grandinatus Gmelin, lectotype in ZMC, specimen figured by Chemnitz, Conchylien Cabinet, vol. 10, pl. 169, fig. 1639, from Palmerston Atoll, Cook Is- lands, 32 x 22.4 mm. Figs. 3-5. Monodonta coronaria Lamarck, MHNG 1096/23, 41 x 27 mm. holotype, (05-452a] May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 71 List of Guide Tabs Cassidae 12-400 For *Tonnidae 13-400 Guide tabs are automatically sent to sub- Binder °Muricidae 13-800 scribers. Additional copies are available at the ° Vasidae 20-400 rate of 10 cents (U.S.) each. The present set of °Harpidae 20-600 16 costs $1.60 (U.S.). Turridae 22-500 List of Issues 00-005 For Pinnidae 53-500 Eat Guide of Contents 00-100 Binder Tridacnidae 62-000 Render Phasianellidae 04-000 3 Index 90-000 l Littorinidae 05-250 Replaced Pages 95-000 Strombidae 09-650 Cypraeidae 10-400 °Text in preparation List of Issues Date of no. First page of Volume 1 Subject Author Publication — pp. looseleaf no. 1, pp. 1-14 Introduction Editors Mar. 31, 1959 14. = 00-003 15- 32. +Vasidae R. T. Abbott . 18 20-403 y, 33-146 Strombus R. T. Abbott Nov. 23, 1960 114 09-831 3; 147-174 — Lambis R. T. Abbott Sept. 28, 1961 38 10-051 4, 175-226 —-Pinnidae J. Rosewater . 52 53-501 2 227-346 Turrinae A. W. B. Powell Mar. 31, 1964 120 22-661 6, 347-396 _Tridacnidae J. Rosewater Apr. 30, 1965 50. 62-003 397-398 Strombus oldi W. K. Emerson . 2 09-881 399-400 — Strombus Listeri T. Okutani 2 10-001 401-402. Miocene Indian Strombus R. T. Abbott 2 09-945 403-406 Turrinae (replacement) A. W. B. Powell 4 22-947 407-408 — Issues and Changes Editors 2 00-005 GG 409-444 ‘Turrinae (concluded) A. W. B. Powell May 15, 1967 36 =. 23-101 445-454 Terebellum Jung and Abbott 7 1009-801 455-456 — Strombus wilsoni R. T. Abbott 2 09-919 8, 457-484 Cypraea (Zoila) Wilson and McComb Dec. 8, 1967 28 = 10-457 485-488 — Index to vol. 1 Editors ° 4 91-001 489-490 — Issues and Changes Editors 2 00-007 Volume 2 no. 9, l- 6 Title pages and Issues Editors Aug 30, 1968 6 00-009 7-202 Cassidae R. T. Abbott ° 196 = 12-409 10, 203-206 Masthead pages Editors Sept. 9, 1969 4 (00-003 207-416 Turriculinae A. W. B. Powell 7 210 23-191 i 417-506 — Littorininae J. Rosewater Nov. 30, 1970 90 = 05-261 12, 507-534 Tectariinae J. Rosewater Jan. 15, 1972 28 05-441 Volume 3 no.13, 1- 40 Drupa Emerson & Cernohorsky May 30, 1973 40 14-801 14 41- 62 Gabrielona R. Robertson 7 22 04-15] 63- 70 Tectarius (replacement) J. Rosewater June 1, 1973 8 05-443a 71- 74 Title page and Issues Editors 4 00-009a [o0-009a] INDO-PACIFIC MOLLUSCA Monographs of the Marine Mollusks of the World with Emphasis on those of the Tropical Western Pacific and Indian Oceans EDITED BY R. TUCKER ABBOTT VOLUME 3 Published by DELAWARE MUSEUM OF NATURAL HISTORY Box 3937, Greenville, Delaware 19807, U.S.A. 74 Title Page Editors Volume 3 a Notice Looseleaf subscribers should place this title page in the first binder immediately after the title page of volume 2. It will be your record of the number of volumes in your looseleaf set. [this sheet issued June 1, 1973, with vol. 3, no. 14] 7 November 27, 1973 5G OS EH Péfett INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae ~~ our LI bes SECTIONAL BRARY DIVISION OF MOLLUSK THE PATELLID LIMPETS OF THE WORLD (PATELLIDAE) by A. W. B. PowrELi Auckland Institute and Museum Auckland, New Zealand Introduction This monograph deals with the Patellidae, one of the five families of limpets that comprise the superfamily Patellacea. The patellid limpets are of littoral and shallow-water occurrence and _al- though widely distributed, cannot be considered cosmopolitan, since they are absent from certain extensive areas, namely, both coasts of North America, the Caribbean, and South America, north of Chile and Patagonia. Although this work is concerned primarily with the Indo-Pacific fauna, it is deemed necessary to extend the scope to world coverage in order to explain the otherwise apparently anomalous dis- tributional patterns. A complication is encountered with the decep- tively similar shells of an allied family, the Ac- maeidae, members of which are easily separable from the Patellidae upon anatomical grounds, but the shell of which usually has no character that can be considered consistently diagnostic; hence, with fossil limpets there is often an element of doubt regarding family allocation. Where patellids are absent, notably along the North West American coast, the acmaeids take over the corresponding littoral, ecological niche, and in so doing, attain shell sizes very large for acmaeids, which usually are of relatively smaller size than patellids. A. striking instance of giant- ism is the Californian Lottia gigantea Gray, 1834, which may reach a length of four inches. Limpets tend to vary greatly in size, shape, sculpture and colour pattern, due to the ecolog- ical factors involved, particularly the relative ex- posure to wave stress and the nature of the sub- stratum. Often, specific limits are apparent only when extensive series from a number of stations are studied. Limpets featured frequently in early concholog- ical works, but many of the species named are difficult to determine with accuracy, since, for the most part, they were based upon crude fig- ures, inadequate descriptions, and with uncertain locality data. Under the heading of cluded in the Patellidae” “Species no longer in- (pp. 84 to 87) 259 Plate 60. Gill structures in the Acmaeidae, Patellidae and Lepetidae. A=Acmaea_ virginea (Miiller); bp = branchial plume, be- hind head. B= Patella vulgata Linnaeus; ecc = gill cordon, complete. = Cellana radians (Gmelin); gei gill cordon, interrupted over head area. D= Nacella (Patinigera) terroris complete; epf= epipodial fringe Patinigera ). k= Lepeta coppingeri (E. A. Smith); no gills; respiration by means of cilia-lined pallial groove. (Fig. A from Fretter and Graham, 1962, p. 120, fig. 73. Fig. E from Eales, 1923, p. 6, fig. 3). (Filhol); (only in gece = gill cordon Nacella and [01-551] e es] 76 Patellidae species, described as Patella are listed, and their present familial location indicated. This list in- cludes species now known to belong to the Ac- maeidae, Lepetidae, Phenacolepatidae, Coccu- linidae, _-Fissurellidae, Stomatellidae, Calyp- traeidae, Capulidae, Hipponicidae, Muricidae, Trimusculidae, Siphonariidae, | Umbraculidae, Ancylidae, and even one considered to be based upon one of the accessory plates of a member of the Pholadidae. Family Patellidae Rafinesque, 1815 The family Patellidae is one of three widespread families——the Acmaeidae Carpenter, 1857, the Patellidae Rafinesque, 1515, and the Lepetidae Dall, 1869, all belonging to the superfamily Patellacea. Two other families, consisting of fossil species only, are placed provisionally in the Patellacea; they are the Metoptomatidae Wenz, 1935, of the middle Silurian to middle Permian, and the Sym- metrocapulidae Wenz, 1938, of the Triassic, Jurassic and possibly Cretaceous. A. W. B. Powell The limpet shell is a simple shield or cap- shaped structure, and is unfortunately a_shell- form that is simulated by molluscs belonging to several other gastropod orders. The one most frequently mistaken for a patellid limpet is Si- pPhonaria, a member of the air-breathing pul- monates, being almost at the top rung of the gastropod ladder of evolution, whereas the Patel- lacea are located down towards the foot of the ladder. In between are the fissurellids, crep- idulids, capulids and umbraculids, all of which have certain members that have limpetlike shells. Limpetlike shells have developed independantly in direct response to ecological necessity, being the shell-form affording the greatest amount of suction area for clinging to a rock surface, cou- pled with a low profile to withstand wave stress. Classification of the patellid limpets, therefore, is dependent upon some knowledge of the animal. Even the allocation of species to either the Ac- maeidae or the Patellidae, the two major families of the Patellacea, in many instances becomes conjectural upon the evidence from the shell alone. Anatomical Outline of the Families and Major Genera of the Patellacea Family Acmaeidae A Leaf-shaped ctenidium only ............... B Ctenidium present, plus gill cordon. Jeans Setdnseceiiocenna asta ntoian acaba eae Acmaea Radula; closely spaced pair of centrals alternating with wider spaced pair of laterals; marginals vestigial or absent. Gill cordon complete ................... Gill cordon interrupted by head Ey enh ot Saree SEEDER EIS ee cere te Scurria sche tuicnsien date Se hain tahoe ae Oe ER: Lottia Family Patellidae C Ctenidium absent; replaced by gill cordon. Radula; 4 or 5 central teeth, median one present, vestigial or absent; lateral large, pluricuspid; marginals 3, weak or vestigial. Gill cordon complete .................. Radula; median central vestigial or absent; alternate pairs of long curved centrals and laterals; marginals 3, weak to vestigial. Gill cordon interrupted by head. Epipodial fringe absent ............ Gill cordon complete. Epipodial fringe present .......... Family Lepetidae D No gills; respiration by cilia-lined pallial groove. Radula; large central, with prominent dentate cusp; no laterals; 2 functional Marcinals ciicsssoemsioriaes Culiigoeeeaiunen ica mend easeenma siete Lepeta November 27, 1973 Biology The embryo of Patella hatches 24 hours after fertilization. The trochophore is about 0.15 mm. in diameter, with a tuft of erect apical cilia and two rows of ciliated cells around the greatest perim- eter of the larva. The cilia beat in clockwise man- ner and rotate the top-shaped larva through the water. Two days after fertilization the larva is transformed into a pretorsional veliger, and both shell and foot appear. Torsion then begins while the larva is free-swimming. During the next stage of about 30 hours the larva both swims and crawls, and torsion is completed when the larva is 3% to 4 days old, and this marks the end of its pelagic life. The velum does not disappear until the snail has been actively crawling for about the third week. About this time the operculum is lost. The shell of the veliger is a dextral coil of scarcely one whorl, and this is soon replaced by a new shell, after which the limpet’s post-larval life con- tinues into the adult. (see Fretter and Graham, 1962, pp. 448-450 for a more detailed account). The English Patella vulgata is a protandrous hermaphrodite with most, if not all, individuals starting life as male but later changing to female at the age of one year or more. Investigations of English populations of vulgata have shown that 90% of the limpets between 16 and 25 mm. in length are male; in those about 40 mm. in length the sexes are about equal; and in those 60 mm. or more in length most are female (see Fretter and Graham, 1962, p. 372). Most patellids feed upon small species of living algae, but some live upon giant kelp, where they scrape away the surface tissue. The rock-dwelling patellids feed with the head end moving method- ically from side to side, while the radula operates like a scythe. Patellids often travel up to four or five feet in search of food, and usually manage to return to their original resting places. This some- times involves re-finding a site previously ex- cavated in the rock that exactly fits the indenta- tions of the shell margin. When rock faces have a slight coating of silt, limpet journeys can be seen quite plainly, and possibly the limpet uses its out- ward track in finding its way home. The age attained by limpets varies greatly according to the species involved, the food poten- tial of the particular habitat, and the zone in which the species occurs. Fretter and Graham (1962, p. 501), quoting Russell (1909), recorded that Patella vulgata from certain established pop- ulations in Scotland attained a length of about 29 mm. in the first year, during which time they INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae = 77 reached sexual maturity. The estimated sizes reached for each of the four succeeding years were respectively 38, 44, 48 and 53 mm. On the other hand, the same species under more favour- able conditions at Plymouth reached a length of 53 mm. by the end of the second year. The esti- mated life span for vulgata is about 15 years. Very large and massive species, such as mexi- cana and kermadecensis, probably live for a much longer time, but size is governed also by the availability of a suitable rock substrate, not un- duly encroached upon by barnacles, corals, or other encrusting marine growths. In the tropical Indo-Pacific it is unusual to find large-sized lim- pets associated with coral reefs, unless there are intrusions of basaltic lava. In general large-sized limpets are more frequent in cool-temperate wa- ters than they are in the tropics. Limpets that live in the upper tidal zone are usually taller than individuals inhabiting the lower tidal zone. Fretter and Graham (1962, p. 501), quoting Orton (1932), correlated these differences with the degree of exposure to desiccation. Lim- pets living near low water mark remain exposed for only a short period by the tide, but high-water limpets are uncovered for the greater part of the day, and this results in a general drying out of the habitat. To prevent desiccation the limpet must hold firmly to the rock for the whole period that it is out of water, and it is suggested that this con- stant application of force by the attachment mus- cles tends to pull in the mantle skirt, which is responsible for new growth around the. shell margin. This produces a taller and narrower shell than that produced by an animal living lower down in the tidal zone. Limpets from very exposed situations often ap- pear to be very different from examples living in more sheltered situations. In the exposed ex- amples, the shell becomes very flat in order to withstand wave stress. As a result of this lowered profile, the apex is located nearer to the anterior end. The colour patterns exhibited by juvenile lim- pets tend to be less variable than those in more mature examples of the same species, and often provide more satisfactory diagnostic criteria. This is especially evident in Cellana strigilis popula-. tions from the southern islands of New Zealand (Powell, 1955, pp. 65-67). Surface erosion of the shell also greatly alters the colour pattern. Thomson (1919, pp. 264-267) described how erosion in Cellana radians often eliminates the transverse “earlii” pattern, but [01-553] 7S Patellidae the pigmentation of the radial ribs survives due to deeper impregnation of the shell substance. With the shrinkage of the animal in senile ex- amples, a thick, unicoloured callus is built up on the inside of the shell, blotting out any macula- tions that may have survived external erosion. The Patellidae are the most successful and the most advanced family of the Patellacea. The change from a simple leaf-shaped ctenidium in the Acmaeidae, to a gill cordon in the Patellidae results in more efficient aeration. Although Scur- ria and Lottia have developed a gill cordon they still retain the acmaeid ctenidium. In the Patel- lidae the ctenidium has entirely disappeared, leaving the gill cordon as the sole means of res- piration. Patellid limpets have become so successful in their chosen littoral environment that in some locations, South Africa in particular, certain spe- cies have become the dominant organisms of sev- eral animal communities. In the “Cochlea zone” of South Africa the species Patella cochlea is so abundant that almost all other forms of animal life are crowded out. A density of 1,300 individ- uals of this limpet to the square yard has been recorded, in so dense a concentration that as many as 40 small individuals were found crowded on top of a single large shell. The radula The radula in the Patellidae is long and narrow, especially in Cellana, in which it sometimes has a length of as much as four times that of the shell. In situ it is concentrated in loose coils on the left hand side when viewed from above. On the other hand, Patella has a much shorter radula that folds back upon itself at the nascent end. The Patella radula consists of a strong or weak or rarely absent median central, flanked by a pair of centrals on either side, followed by a large pRiricuspid lateral, and finally, three weak, slen- der, functionless marginals. The latter may be fused into a single plate. Well-developed cusps, capped with a dark stain containing magnetite, are present on the multiple centrals and the pluri- cuspid laterals, but cusps on the remaining teeth are small to vestigial, and colourless. In Cellana and Nacella the radula differs from that of Patella, in that the functional teeth are a pair of long, large, centrals, closely-spaced, on either side of a vestigial plate, which also may be absent, alternating with a wider-spaced pair of similar, well-developed laterals; the functionless marginals are as in Patella. A. W. B. Powell In Patella the multiple centrals and the large pluricuspid lateral have relatively short recurved cusps, but in Cellana and Nacella, the alternating pairs of centrals and laterals are very long and project arcuately upward, almost at right angles to the base. These long, strongly upcurved teeth present difficulty in slide preparation, for they are easily pressed at varying angles in mounting, and thus may assume very different shapes. A satis- factory solution to this problem is in the use of cavity slides, that bridge the radula across, so that the teeth assume their normal upright posi- tion over the cavity, but are pressed sideways, beyond the limits of the cavity, thus giving details of denticles or indentations along the sides of the teeth. Many writers have endeavoured to use the length of the radula in relation to the length of the shell for separating three assumed closely allied English Patella. The range of the means arrived at by Fretter and Graham (1962, p. 495) is tabulated below. Plate 61. Fig. 1. Patella vulgata Linnaeus; England. Radula. Fig. 2. Patella caerulea Linnaeus; Trieste. Radula, from Thiele, in Troschel and Thiele, 1891, pl. 25, fig. 18. [01-554] November 27, 1973 vulgata intermedia aspera Length of radula Sis 1.60-2.10 Length of shell 1.05-1.15 Unfortunately, in that work, there is no precise indication of the actual identity of the species termed Patella intermedia, there being four dif- ferent usages of that combination by four differ- ent authors. Brian and Owen (1952, pp. 241-249) provided a useful table, giving the valid name equivalents for the nomenclature used in papers on European Patellidae, published up until 1948 (see under heading of Patella intermedia Auct.). Following is the summary of Brian and Owen’s conclusions. “Patella vulgata L. were collected from high- and low-water levels on five different beaches and the lengths of the shells and radula measured. In all five localities the mean shell length was less and the mean radula length greater at the higher level, but the difference was not consistently sig- nificant.” “The complexity arose from the mergence of two conflicting tendencies: one, intralevel, a posi- tive regression of shell-length and radula. While the former is no doubt a growth phenomenon, the latter is probably an environmental effect—at higher level exposure to desiccation and conse- quently, prolonged adherence to the substrate, caused a smaller shell base; reduced feeding time caused a longer radula. These factors may have resulted in confusion when comparing the values of the radula fractions of species of Patella.” Lowenstam (1962) has shown that the radular teeth in the limpets, Acmaea, Lottia and Patella, are capped with goethite, a dark opaque mineral of high iron content. This gives the dark-stained primary cusps of the patellacea a hardness of al- most 5 on the Moho hardness scale, whereas the radular hardness in the Littorinidae, the Fissurelli- dae and certain trochoids is only between 2 and 3. This hardness factor in tho radula enables lim- pets to erode limestone and other rocks of com- parable hardness, as well as the external surface of other shells. Limpets frequently excavate deep depressions in the rock to form a base of attach- ment, and are known to return to their own par- ticular site after a nightly foraging excursion. Lowenstam (1962a) also suggested that since, in the case of chitons, the dark stained denticle caps show the presence of magnetite, then it is possible that these magnetised teeth may serve as a guidance system for the so-called homing in- stinct of both chitons and limpets. INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae 79 Geographical Distribution of the Patellacea The family Patellidae has achieved a very wide distribution, extending from near the Arctic Circle to the Antarctic Circle, the western Atlantic, Med- iterranean, West and South Africa, the Indo- Pacific to as far north as northern Japan, east- ward to the Hawaiian Chain and the Island of Juan Fernandez, and southward to Australia, New Zealand, the subantarctic of the southern end of South America, the islands of the Southern Ocean, and even a few locations adjacent to the Antarctic Continent. Strangely, there are no patellids on either the east or the west coasts of North America, the Caribbean and most of South America. The only exceptions are the giant Patella (Ancistromesus) mexicana from tropical Central America, obvi- ously derived from the Indo-Pacific when the former Tethys Sea was an open waterway around the perimeter of the globe, and the cold water Nacella-Patinigera complex that is strongly repre- sented in southern Patagonia and Chile, and from there spread eastward to the islands of the South- ern Ocean. The Recent distributional patterns for each of the three families of the Patellacea are outlined below (see plate 62). Acmaeidae: The typical genus, Acmaea, is of worldwide distribution, but has its greatest devel- opment in western North America, where it takes the place of the Patellidae, members of which are absent from that area. Two other genera of the Acmaeidae, Scurria and Lottia, are apparent fore- runners of the Patellidae, since they have devel- oped pallial gills, additional to the single gill- plume of typical Acmaea. Their present range is along the west coast of both North and South America. Patellidae: Typical Patella is well-represented along the western coast of Europe, from the Lofo- ten Islands to Great Britain, down to Spain, the Mediterranean, the West African mainland and off-shore islands, then southward to South Africa, where typical Patella becomes more or less merged into a composite local fauna of cold and Wwarm-water subgenera. From there the genus continues, in subgeneric form, across to the warmer water island groups of the Indo-Pacific, extending northward to Japan, eastward to the west coast of tropical Central America, and south- ward to the Kermadec Islands and the temperate waters of southern Australia and Tasmania. The genus Cellana is confined for the most part to the warm and cool temperate waters of the [01-555] SO. Patellidae A. W. B. Powell Indo-Pacific, but in the New Zealand area it ac- tually extends southward into subantarctic waters. The farthest westward reached by Cellana is the coast of Natal, the farthest eastward the island of Juan Fernandez, off the coast of Chile, and the farthest northward, Japan. A third major genus in this family is Nacella, containing the important subgenus Patinigera. These are truly cold-water limpets, the greater part of their range being subantarctic, but extend- ing to the Antarctic by way of the Scotia Arc, and also ranging northward up the Chilean coast to at least Valparaiso, assisted in this by the upwelling of cold water along that coast. The present con- centration of the genus is in the Magellanic area, and from there it spreads eastward, assisted in this by the prevailing West Wind Drift. Many of the species live upon the large kelps which pro- vide an effective means of chance dispersal when quantities of the weed are wrenched free and drift before wind and current. The farthest east- ward that this genus has established itself is Ker- guelen Island, and for the subgenus Campbell Island in the New Zealand southern islands. The genus Nacella and its subgenus Patinigera have a distinctive epipodial fringe, not found so far in any other genus of the Patellacea. The European Tertiary fossils attributed to Nacella probably belong to other genera. Lepetidae: This family consists of rather small featureless white limpets, mainly from the deeper waters of the Arctic Ocean, the north Atlantic, the north Pacific, Mediterranean, Patagonia and Antarctica. They are rather specialised, but not necessarily highly advanced. There are no gills, and respiration takes place through a cilia-lined pallial groove. The presence of a large, broad- based central tooth, with a conspicuous, broadly- triangular, dentate cusp, no laterals, but a pair of functional marginals, are radular characters not found in the other two living families of the Patel- lacea. Fossil Occurrences of the Patellidae Although a considerable number of fossil so- called Patella species are encountered in liter- ature, especially those from European Tertiary localities, few of them can be assigned with cer- tainty to that genus. The problem faced by workers with Recent species of the Patellacea, that of distinguishing between the Acmaeidae and the Patellidae when the animal is unknown, is even more a matter of conjecture when fossil species are under consideration, especially with those from the older formations, that have shell features unlike those of living species. The earliest species of the Patellidae that can be generically identified with some degree of con- Ft NORTHERN LIMIT of PATELLA ¢ [P.(SCUTELLASTRA er yee ye HELCION & a en PATINASTRA * a ee of a / NORTHERN LIMIT OF NACELLA and PATINIGERA o / ot o ——: pain =" [NACELLA and PATINIGERA + T 140 1090 Plate 62. Geographical distribution of the Patellidae. Note the almost entire absence of the family from North America, and most of South America, with the exception of one species, Ancistromesus mexicana, which occurs along the west coast of Central America, and Nacella, with its subgenus Patini- gera, in the southern part of South America, from where it has drifted eastward over much of the Subantarctic and in some areas of the Antarctic. [01-556] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae $1 Plate 63. European and South Africa Patella Figs. 1-3. Patella vulgata Linnaeus, 1758. Figs. 1, 2. Caldy Island, South Wales. Fig. 3. [fracombe, England. Fig. 4. Patella aspera Roding, 1798. Caldy Island, South Wales. Figs. 5-7. Patella caerulea’ Linnaeus, 1758. Isle of Capri, Italy. Fig. $8. Patella) caerulea Linnaeus, 1758 (forma crenata Gmelin, 1791). Madeira. [01-557] Fig. 9. Patella (Cymbula) compressa Linnaeus, 1758. Kom- metje, Cape Peninsula, South Africa. Lives on large kelp; always sideways compressed, Figs. 10, 11. Patella (Cymbula) miniata Born, 1778. Fig. 10. False Bay, South Africa. Fig. 11. Port Elizabeth, South Africa; beach shells bleach to bright pink. Figs. 12-14. Patella (Olana) cochlear Born, 1778. Sea Point, South Africa. Anterior end always constricted like a spout. $2 Patellidae A. W. B. Powell fidence are Cellana carpentariana Skwarko, 1966, from the late Neocomian lower Cretaceous of Northern Territory, Australia, and Patella (Scutel- lastra) cooperi (Powell, 1938) from the Otaian Stage, lower Miocene of Motuihi Island, Auck- land, New Zealand. The above two records show at least that the separation of Patella and Cellana is of long stand- ing, and coupled with the radular divergence, justifies Thiele’s division of the family into two subfamilies, the Patellinae Rafinesque, 1815, and the Nacellinae Thiele, 1929. Doubtful species of Patellidae Patella ? amuritica Wilckens, 1922 Range—New Zealand, Amuri Bluff, upper Cre- taceous. Remarks—This species is based upon a very damaged and incomplete shell only 5 mm. in length. The whole of the apical area is missing and only a ring of shelly material remains. Even Wilckens expressed doubt as to whether his gen- eric determination was correct. Synonymy— 1922 Patella ? amuritica Wilckens, N. Z. Geol. Surv. Pal. Bull. no. 9, p. 5, pl. 1, fig. 8. Patella guineensis Dunker, 1853 Remarks—The_ present writer has insufficient West African material to evaluate Dunker’s spe- cies, the name of which is preoccupied by Patella guineensis Gmelin, 1791. Synonymy— 1853 Patella guineensis Dunker, Ind. Moll. Guin. Infer., p. 49, pl. 7, figs. 1-3; 19-21. Loanda, Guinea, West Africa [Angola]. Cellana jutsoni Chapman and Crespin, 1934 Remarks—This species, from the lower Miocene Plantagenet Beds of Albany, Western Australia, is very doubtfully patellid. The presence of an “obscure ridge, extending from the apex to the posterior margin” suggests the fissurellid genus Tugali or something akin to it. Synonymy— 1934 “Cellana” jutsoni Chapman & Crespin, Journ. Roy. Soc. West. Aust., vol. 20, p. 122, pl. 11, fig. 28. Patella aspera Roding, 1798 (Pl. 63, fig. 4; pl. 68, figs. 3, 4) Range—British Isles and Atlantic coast of France. Remarks—This “species” is not always readily distinguished from vulgata. Typically it is more elongated than vulgata, depressed, with the apex nearer to the anterior end, and the primary ribs are stronger and sharper, resulting in a more def- initely corrugated margin. Other differences are that the interior is porcellanous whitish, with the head scar pale orange, radial colour lines are sub- obsolete to obsolete, and the colour of the foot of the animal is cream to orange, as opposed to grey-green in vulgata. Measurements (mm.)— length width height 53.0 42.0 21.0 Caldy Id., South Wales 47.0 35.5 20.0 Caldy Id., South Wales Synonymy— 1798 Patella aspera Roding, Mus. Bolten., vol. 2, p. 10 (refers to Favanne, pl. 2, f. G). 1819 Patella aspera Lamarck, Anim. sans Vert., vol. 6, p. 327 (refers to Favanne, pl. 2, f. G). 1844 Patella athletica Bean, in Thorpe, Brit. Mar. Conch., p. 264, fig. 101). 1968 Patella aspera Lam., McMillan, Brit. Shells, Warne & Co. Ltd., London. New York, p. 25, pl. 1, figs, 1, 3, 4. Patella depressa Pennant, 1777 Range—South coast of England, Channel Islands and Atlantic coast of France. Remarks—This is the small, very depressed, Siphonaria-like species, or form of vulgata, of which Patella vulgata var. intermedia Jeffreys, 1865 is a synonym. Jeffreys described his variety intermedia as “Shell rather smaller, flatter, and oval, with finer ribs, and an orange crown; inside golden-yellow or. tinged with flesh colour (oc- casionally cream colour) in the centre, and beau- tifully rayed toward the margin.” Forbes also re- marked that the animal is black or dark-coloured. Despite the detailed studies of both Fischer- Piette and R. G. Evans, the taxonomic status of both aspera and depressa in relation to vulgata is still uncertain. Evans endeavoured to separate the three as full species, upon minute differences in the pluricuspid radula teeth, coupled with the varying lengths of the radula for each. In dealing with populations from the south of England Evans admitted, that at the Isle of Wight, intermediate forms were common but then remarked that to the westward along the south coast three species form discontinuous entities. [01-558] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae 83 Synonymy— 1777 Patella depressa Pennant, Brit. Zool., vol. 4, p. 124, pl. 89, fig. 146. 1865 Patella, vulgata var. intermedia Jeffreys, Brit. Conch. vol. 3, p. 237. 1923 Patella depressa Pennant, Tomlin, Journ. Conch., vol. 17, p. 34. 1935 Patella spp. Fischer-Piette, Systematique et biogeo- graphie-Les Patelles d’Europe et d'Afrique du Nord, Journ. Conchyl., vol. 79, pp. 5-66. 1952 Patella depressa Pennant, Evans, Proc. Zool. Soc., Lond., pp. 357-376. Patella electrina Reeve, 1854 Remarks—This shell, described as coming from Australia, is unlike any species known from that area. Examination of the type specimen in the British Museum (Natural History) revealed a Patella of the caerulea series, very like the Canary Islands Patella lowei d’Orbigny, so far as one can judge from a single example. Synonymy— 1854 Patella electrina Reeve, Conch. Iconica, pl. 22, fig. 55a, b. Patella intermedia Many writers have attempted to give taxonomic status to the forms of both vulgata and caerulea by providing detailed studies of the radulae, par- ticularly its length in relation to that of the shell (see also, under the heading “Radula,” in the in- troduction), in small differences in the pluricuspid laterals, and also in the colour of the foot of the animal. Unfortunately with many of these papers it is difficult to correlate the results with the species or forms investigated, due to a common fault of many anatomists of placing little or no importance upon the characters of the shell, and seldom illus- trating the relevant shells. Another source of confusion is in the nomen- clature employed, specific names being frequently cited without their respective authority and date. A name that is quoted frequently, and around which much useful data is associated is “inter- media,” but which patellid of that name is in- tended?—that of Knapp, 1557, Jeffreys, 1565, or Bucquoy, Dautzenberg and Dollfus, 1852? Brian and Owen (1952) endeavoured to rectify this confusion by concluding that P. intermedia Jeffreys, 1865, as used by Fischer-Piette (1935, 1938 and 1948), Eslick (1940) and Orton (1946) was Patella depressa Pennant, 1777. They also concluded that P. athletica Bean, 1844, as used in the Plymouth Marine Fauna (1931) and by Winckworth (1932) and Evans (1947) and P. de- pressa, as used by Fischer-Piette (1935), Eslick (1940) and Orton (1946) were all Patella aspera Lamarck (now aspera Roding, 1798). The earliest use of the name intermedia, in association with Patella, is that of Knapp, in Murray, 1857, proposed for a Channel Islands shell, that appears to be a form of Patella vulgata Linnaeus, 1758. Patella lineata Lamarck, 1819 Remarks—This shell, from unknown locality, was referred to Helcioniscus (now-Cellana) by Pilsbry, but Delessert’s figures suggest a species of the Acmaeidae. Pilsbry’s translation of Lam- arck’s description follows: Description—“Shell oval, convex, buff-brown, painted with 10-12 yellow lines; excessively nu- merous longitudinal close striae; vertex acute, buff. Length exceeding one inch.” - 27 mm. (Mer- mod). Synonymy— 1819 Patella lineata Lamarck, Anim. sans vert., vol. 6 (1), p. 331; Patella lineata Delessert, Rec. de Coq., pl. 23, fig. 6. 1891 Helcioniscus lineatus Lamarck, Pilsbry, Man. of Conch., vol. 13, p. 153, pl. 73, figs. 85-87. 1950 Patella lineata Lamarck, Mermod, Revue Suisse de Zool., vol. 57, no. 34, p. 694 (remarks on the type). ? Patella nelsonensis Trechmann, 1918 Range—Nelson, New Zealand, lower conglom- erates, lower slopes of range, upper end. of Eighty-eight Valley, Kaihikuan Stage, middle Tri- assic. Remarks—The holotype and two paratypes are in the New Zealand Geological Survey, Welling- ton, but the preservation is not good enough to show muscle scars, hence it cannot be deter- mined, on the present material, whether the apex is directed anteriorly or posteriorly. The species remains a doubtful member of the Patellacea. Synonymy— 1918 Patella (?) nelsonensis Trechmann, Quart. J. Geol. Soc., vol. 73, pt. 3, p. 185, pl. 18, figs. S a, b. 1953 Patella nelsonensis Trechmann, Marwick, N. Z. Geol. Surv. Pal. Bull. no. 21, p. 74, pl. 7, fig. 3. Patella reussi K. Martin, 1879 Remarks—This species from the Miocene of Java could be fissurellid. The figure shows a shell em- bedded in matrix, and in consequence the interior [01-559] $4. Patellidae of the shell, with its muscle impressions, is un- known. Synonymy— 1879 Patella reussi K. Martin, Die Tert. auf Java, Leiden, p. 87, pl. 12, fig. 9. Patella spectabilis Dunker, 1853 Remarks—The_ description and_ figures of Dunker’s species suggest a form of Patella lowei d’Orbigny, 1839. However, Dunker’s name is pre- occupied by Patella spectabilis Gmelin, 1791. Synonymy— 1853 Patella spectabilis Dunker, Ind. Moll. Guin. Infer., p. 39, pl. 6, figs. 7-9. Loanda, Guinea, West Africa [Angola]. Species no longer included in the Patellidae The following species, originally referred to the Patellidae, are now known to belong to other families. This list includes only species that the writer has been able to evaluate, either per- sonally, or upon the authority of subsequent revisers. There remains a considerable number of patellid names yet to investigate, but since many of these have been inadequately described, sel- dom figured, and often without locality data, it is probable that most, eventually, will have to be considered indeterminate. Species prefixed by an asterisk (*) are the sub- ject of a note at the conclusion of this section. The writer has compiled a manuscript list of over 1,250 patellid names encountered in the preparation of the present work, but it is withheld from publication at this stage, since it cannot be considered reasonably complete, without a more thorough appraisal of European species, the fossil ones in particular. A dagger (+) precedes fossil species in this list. achates Reeve, 1855, Patella ............ccccccccseeeeeseeneeeeee Acmaeidae acinaces Lea, 1846, Patella .........0...ccc cece eee ceeeeeeee ees ? Pholadidae aculeata Gmelin, 1791, Patella 2.0.0.0... ccsseseens Caly ptraeidae adunca Perry, 1811, Patella... eee Fissurellidae aenigmatica Mabille, 1895, Patella 00.0000... Acmaeidae aeruginosa Middendorff, 1847, Patella (Acmaea) ... Acmaeidae afra Gmelin, 1791,: Patellas wsisssccsiuscsctnaantessaccrantvens Trimusculidae albescens Anton, 1839, Patella ...........cccccceeceeeeeeeeees Acmaeidae albicosta C. B. Adams, 1855, Patella ..............:008 Acmaeidae albicostata Reeve, 1855, Patella .....0.....0..cccccceeeeee ees Acmaeidae alticostata Angas, 1865, Patella w.........ccccccceeecseeee tees Acmaeidae alveus ‘Conrad, 183i... Patel adits nnconaaiisaiennecenanats: Acmaeidae ambigua Wood, ISIS, Patella ...........0.ceeeeeeeeeees Fissurellidae amoena, Say; 1822, Patella siiisisscissssccanctecsensedesasesdess Acmaeidae ancyloides E. Forbes, 1840, Patella oo... Lepetidae ancyloides Middendorff, 1847, Patella (Acmaea) ... Acmaeidae angulata Wood, 1828, Patella ............cccceeeeeeeee Calyptraeidae angusta Gmelin, 1791, Patella oo... cece Fissurellidae A. W. B. Powell antillarum Philippi, 1849, Patella (Acmaea) ............ Acmaeidae antiquata Linnaeus, 1767, Patella ...................0 Hipponicidae apertura Montagu, 1803, Patella .......00000.0.....008 Fissurellidae araneosa Gould, 1848, Patella ......00.ccccceceeeeeeeeeeeees Acmaeidae araneosa Reeve, 1855, Patella .......0.0..cceceeceee cece eee Acmaeidae araucana d’Orbigny, 1841, Patella .....000...00...c0 Acmaeidae asmi Middendorff, 1847, Patella ......0.0...0.00ccccccee Acmaeidae atricapilla Dillwyn, 1817, Patella oo... Fissurellidae auricula Gmelin, 1791, Patella 0.0.00... Calyptraeidae auricula W. Wood, 1828, Patella. .........0.....:c0cee Stomatellidae australis Lamarck, 1819, Patella ...0......ceees Hipponicidae avellana Gmelin, 1791, Patella .........0000...cccccceeeee Fissurellidae axiaerata Verco, 1912, Patella ............ccccccsscsssseeeeeees Acmaeidae balanoides Reeve, 1855, Patella .....0...000.cccceeee Acmaeidae barbadensis Gmelin, 1791, Patella ....000.....0c.ccce Fissurellidae biradiata Reeve, 1855, Patella ........0.00ceccceeeeeeeees Acmaeidae borneensis Reeve, 1855, Patella ..........00....ccccceceeeeee Acmaeidae borniana Helbling, 1779, Patella .................0000cceeeees Acmaeidae caeca Muller, 1776, Patella ........0...cccccccecceeseeeeesseees Lepetidae calamus Crosse & Fischer, 1864, Patella ............00... Acmaeidae callosa Hombron & Jacquinot, 1841, Patella .......... Acmaeidae campaniformis Blainville, 1825, Patella ............ ? Siphonariidae campbelli Filhol, 1880, Patella ....00...00...cccee Acmaeidae cancellata Gmelin, 1791, Patella ......00.00.ccceee Acmaeidae candida Couthouy, 1838, Patella 0.0.0.0. Lepetidae cantharus Reeve, 1855, Patella ........cccccccccceeeeeeeceeeeee Acmaeidae cassida Dillwyn, 1817, Patella oo... Hipponicidae casta Carpenter, 1866, Nacella .............cccccceecceeeeees Acmaeidae ceciliana d’Orbigny, 1841, Patella oo... Acmaeidae cerea Mller, 1842, Patella .......0..cccccccccssesseeeeneeees Lepetidae chilensis Blainville, 1825, Patella .....0..00..000. ? Siphonariidae chinensis Linnaeus, 1758, Patella ......000...00..0..... Caly ptraeidae cimeliata Reeve, 1855, Patella ....0..0.0000000 eee Acmaeidae cinis Reeve, 1854, Patella .........0.0.ccccccccessecseseseeeeeeeees Acmaeidae cinnamomea Gould, 1846, Patella ................ Phenacolepadidae clealandi J. Sowerby, 1822, Patella .....0....cce Acmaeidae clypeus T. Brown, 1827, Patella .......00.c.cccccceeeeeeeee Acmaeidae cochleata Dillwyn: 1807, oc.ccccssssieag cose esnintessscaecess Capulidae coffea Reeve, 1855, Patella ............c0cccccccessssseeeeeeeeees Acmaeidae compressiuscula Karsten, 1849, Patella ..........0....... Acmaeidae concentrica Middendorff, 1851, Patella ..............000.. Lepetidae concepsionis Lesson, 1831, Patella ............. Acmaeidae concinna Lischke, 1870, Patella ..................0000ce cee Acmaeidae conica Defrance, 1825, Patella ...............cccccceeeeeceeees Acmaeidae conica Gould, 1846, Patella ........00cccccccceeeeeeeee Acmaeidae conulus Dunker, 1882, Patella .........cccccccccceeecceceeeeeee Acmaeidae corrugata Reeve, 1854, Patella ...........ccccccecceece sence Acmaeidae craniolaris Roding, 1798, Patella oo... eee Fissurellidae crebrestriata Verco, 1904, Nacella .....0...000ccccceeeee Acmaeidae crepidula Linnaeus, 1764, Patella .....0..00..00.006. Calyptraeidae cruciata Linnaeus, 1758, Patella ....0.0.000000.cceeee Acmaeidae crystallina W. Wood, 1828, Patella ...............0000 Fissurellidae cubensis Reeve, 1855, Patella ..........c0cccccccecseeeeeeeees Acmaeidae cumingii Reeve, 1854, Patella ..........0..0..ee Acmaeidae diaphana Reeve, 1854, Patella .........c0cccecccsseceees Acmaeidae dichotoma Anton, 1839, Patella ......0000....cccceeeeeeeees Acmaeidae digitale RGding, 1798, Patella .......0.0ccccccecceeeeeeeeees Acmaeidae discors Philippi, 1849, Patella (Acmaea) ................ Acmaeidae elegans Philippi, 1849, Patella ....0..0000.0cccee Acmaeidae emarginuloides Philippi, 1868, Patella .........00...0.0..... Lepetidae equestris Linnaeus, 1758, Patella... Calyptraeidae exilis Philippi, 1849, Patella ......0..00. 0c ccecceeceeeee Acmaeidae fenestrata Reeve, 1855, Patella ......0000.cccccccceeeeereee Acmaeidae fimbriata Gould, 1846, Patella ...0.00000....cccceeceeeeeeee Acmaeidae fissura Linnaeus, 1758, Patella .............0.000cc cece Fissurellidae fissurata Dillwyn, 1817, Patella 0.0.0... Fissurellidae fissurella O. F. Miiller, 1776, Patella ........0.0c0ccccce. Fissurellidae floccata Reeve, 1855, Patella .......00ccccccccceeeeeeeeeeeees Acmaeidae fluviatilis Gmelin, 1791, Patella ......0.0ccccccceceeeeeee Ancylidae forbesii J. Smith, 1839, Patella oo... cece Lepetidae fornicata Linnaeus, 1758, Patella oo... Calyptraeidae fulva O. F. Miiller, 1776, Patella ...0.0..0ccccceeeee Lepetidae fungoides Réding, 1798, Patella v.00 Acmaeidae [01-560] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae 85 Plate 64. South African Patella Figs. 1-3. Patella (Patellona) granatina Linnaeus, 1758. Sea Figs. 7-9. Patella (Patellona) oculus Born, 1778. Buffel’s Point, South Africa. Bay, Cape Peninsula, South Africa. Figs. 4-6. Patella (Patellidea) granularis Linnaeus, 1758. Figs. 10-11. Patella concolor Krauss, 1848. Near Durban, Sea Point, South Africa. Natal. [01-561] S86 Patellidae A. W. B. Powell galathea Lamarck, 1819, Patella ........00........ Phenacolepadidae goreensis Gmelin, 1791, Patella oo... Calyptraeidae graeca Linnaeus, 1758, Patella ...........000.....cccceeeeeees Fissurellidae grammia Philippi, 1847, Patella... Acmaeidae granostriata Schrenck, 1867, Patella .........0.....0000 Acmaeidae granulata Philippi, 1848, Patella... Acmaeidae grisea Gmelin, 1791, Patella... Siphonariidae grisea Réding, 1798, Patella ...........0..00cccceeeceeceeeeeees Fissurellidae haliotoidea Réding, 1798, Patella ...........0....6. Calyptraeidae hepatica Pritchard & Gatliff, 1903, Patella ............. Acmaeidae heptagona Blainville, 1825, Patella .........0.......08 Acmaeidae heroldi Dunker, 1882, Patella .................ccccccesceeeeees Acmaeidae hiantula W. Wood, 1828, Patella ....0.............00ccce Fissurellidae °hochstetteri K. Martin, 1879, Patella) .................. Fissurellidae incisa Dillwyn, 1817, Patella 20.0000... Fissurellidae inconspicua Gray, 1843, Patella ..........0.. Acmaeidae indica W. Wood, 1828, Patella .................ccceee Umbraculidae inradiata Reeve, 1855, Patella ..............00...ccceeeeeeeee ? Acmaeidae insessa Hinds, 1842, Patella 0.0.0... eee eee Acmaeidae insignis Menke, 1843, Patella ...........0.......cccceccecee eee Acmaeidae instabilis Gould, 1846, Patella ..........0000000.ccc cece Acmaeidae intertexta ROding, 1798, Patella ......000.00..cc eens Acmaeidae jacksoniensis Reeve, 1855, Patella ......0.....cceeee Acmaeidae jamaicensis Gmelin, 1791, Patella p. 3715 w........... Acmaeidae jamaicensis Gmelin, 1791, Patella, p. 3730 ............ Fissurellidae javanica Lamarck, 1819, Patella ................c28. Siphonariidae kochi Philippi, 1549, Patella oo... eee Acmaeidae laciniata Reeve, 1855, Patella ........0......0ccccceeee eee Acmaeidae laciniosa Linnaeus, 1758, Patella ..............ce Siphonariidae lacunosa Reeve, 1855, Patella ................000ccccee eects Acmaeidae lacustris Linnaeus, 1758, Patella ............0..cc eee Ancylidae lamanonii Schrenck, 1867, Patella ..................000:0005 Acmaeidae lanx Reeve, 1855, Patella ............cccccccccccsssseeeeeeeeeeeees Acmaeidae laqueare W. Wood, 1828, Patella ...........:ce ? Acmaeidae latistrigata Angas, 1865, Patella 0... ee Acmaeidae lentiginosa Reeve, 1855, Patella ..............::c0ccceeeeeeeees Acmaeidae lepas Gmelin, 1791, Patella ............0.05. Muricidae (Thaidinae) leucophaea Philippi, 1849, Patella .......000....... Acmaeidae leucopleura Gmelin, 1791, Patella ................0.00 Acmaeidae leucopleura Reeve, 1855, Patella .........0.. cee Acmaeidae lima Reeve, 1855, Patella .............00cccceececeeeeeeeee seen es Acmaeidae limbata Réding, 1798, Patella ...........ceeees Fissurellidae lineata Philippi, 1849, Patella... Acmaeidae *luchuana Pilsbry, 1901, Patella... Acmaeidae luctuosa Hombron & Jacquinot, 1841, Patella ........ Acmaeidae lutea Linnaeus, 1758, Patella ................eee Stomatellidae macroschisma Lightfoot, 1786, Patella .................. Fissurellidae macroschisma Dillwyn, 1817, Patella ..........0....0.... Fissurellidae mamillata (Nuttall) Reeve, 1855, Patella ................ Acmaeidae mauritiana Pilsbry, 1891, Helcioniscus .................5 Acmaeidae melanoleuca Gmelin, 1791, Patella .....................:008 Acmaeidae melanoleuca Reeve, 1855, Patella 0.0.0.0... Acmaeidae merceauxi Deshayes, 1861, Patella ..............:ee Acmaeidae minima Gmelin, 1791, Patella ......0..........c0cccceeeee ees ? Acmaeidae mitella R6ding, 1798, Patella .................cccceeeeeeeeeees Fissurellidae mixta Reeve, 1855, Patella ............ccceeeeeeeeeeeeeeeeeeees Acmaeidae miilleri Dunker, 1875, Patella (Tectura) ................. Acmaeidae muricata Brocchi, 1814, Patella .................e. Calyptraeidae mytiliformis Gmelin, 1791, Patella oo... Calyptraeidae navicula Reeve, 1854, Patella oo... ccc eeeees Acmaeidae neptuni Dillwyn, 1817, Patella 00... Calyptraeidae °nigrosulcata Reeve, 1855, Patella RE Rem Acmaeidae nimbosa Linnaeus, 1758, Patella .................:0 Fissurellidae noachina Linnaeus, 1771. Patella .....0..0...............0- Fissurellidae nodosa Born, 1778, Patella ...........60ccccceceeeeeeeeeeeeeeees Fissurellidae notata Linnaeus, 1758, Patella .....0.......0..ccceeeeeeeeeee Fissurellidae nubecula Linnaeus, 1758, Patella .....................000 Fissurellidae nummularis R6ding, 1798, Patella ..... .... Fissurellidae nuttalliana Reeve, 1855, Patella ...............ceceeeeeee Acmaeidae obliquata Koenen, 1892, Patella (Acmaea) ....... ? Cocculinidae obscura Hombron & Jackquinot, 1541, Patella ...... Acmaeidae occidentalis Reeve, 1855, Patella ...........c.ccccceeeceeeees Acmaeidae octoradiata Gmelin, 1791, Patella Fissurellidae onychites Menke, 1843, Patella ..........0...0cccccceeeseees Acmaeidae topea Reeve, 1854, Patella... eeeeeeeees Acmaeidae oregona Nuttall, 1839, Patella .......0000.00ee .. Acmaeidae pallescens Philippi, 1849, Patella (Acmaea) ............ Acmaeidae tpallida Gould, 1859, Patella ....0...cccceeeee ... Acmaeidae papillaris Réding, 1798, Patella .............00.ccceeee Acmaeidae parasitica d’Orbigny, 1841, Patella ............. . Acmaeidae parva da Costa, 1778, Patella .........0.cccceee ... Acmaeidae patina Eschscholtz, 1847, Patella (Acmaea) ............ Acmaeidae pectinata Linnaeus, 1758, Patella .........000..0..00.. Siphonariidae pediculus Philippi, 1846, Patella .......00.000.00.cccce Acmaeidae peltoides Carpenter, 1864, Nacella. .... .... Siphonariidae penicillata Reeve, 1855, Patella ........0..0.ceeee Acmaeidae perforata Gmelin, 1791, Patella .........0.c eee Fissurellidae persona Eschscholtz, 1847, Patella (Acmaea) ......... Acmaeidae personata T. Martyn, 1788, Patella (non binom.) Fissurellidae perversa Gmelin, 1791, Patella... Umbraculidae personoides Middendorff, 1849, Patella (Acmaea) Acmaeidae peziza W. Wood, 1828, Patella ............ ees Calyptraeidae phyrozonias Gmelin, 1791, Patella oo... Fissurellidae picta Gmelin, 1791, Patella .......ccceeeeeeereees Fissurellidae pileolus Dillwyn, 1817, Patella wo... cee Fissurellidae pileolus Middendorff, 1849, Patella (Acmaea) ....... Acmaeidae plana Philippi, 1849, Patella 00... eee Acmaeidae plana Reeve, 1855, Patella ..............0cccccccceceeeeeeeeeeens Acmaeidae poculum W. Wood, 1828, Patella ..............0:00 Calyptraeidae pretrei d’Orbigny, 1841, Patella ....00...00.cc eee Acmaeidae tprofunda Deshayes, 1863, Patella .............0.:c Acmaeidae punctatissima Philippi, 1849, Patella... Acmaeidae punctulata Gmelin, 1791, Patella ................ccccceeeeees Acmaeidae puncturata Lamarck, 1819, Patella 0.0... Acmaeidae pustula Gmelin, 1791, Patella .........00.0000c ccc Fissurellidae pustulata Helbling, 1779, Patella 0... eee Acmaeidae pygmaea Dunker, 1882, Patella .............00000cc eee Acmaeidae rosea Gmelin, 1791, Patella ............000ccc cece ceeeeeeeee eee Fissurellidae rosea Dall, 1872, Nacella ? wc... ccccccecccceesceeeesceeneeeees Acmaeidae roseoradiata Verco, 1912, Nacella crebrestriata ..... Acmaeidae rubella O. Fabricius, 1780, Patella .........000......cceeee Acmaeidae rubeola R6ding, 1798, Patella ........000000.....0ceeeee Fissurellidae rudis R6ding, 1798, Patella ...............0ccccceseeeeseeeeeees Fissurellidae rugosa Réding, 1798, Patella 0.0.0.0... Capulidae saccharina Linnaeus, 1758, Patella... Acmaeidae scabra Gould, 1846, Patella (Lottia) ......000000.0....... Acmaeidae scabra Reeve, 1855, Patella ..........eceeeeeeseeeeeeeeeeees Acmaeidae scapula Martyn, 1789, Patella (non binom.) ............ Aplysiidae schrenckii Lischke, 1868, Patella ........00......00:cee Acmaeidae scurra Lesson, 1831, Patella ...........cccccceeeeeeeeeeees Acmaeidae scutellata W. Wood, 1828, Patella oo... Calyptraeidae senilis R6ding, 1798, Patella... Umbraculidae sinensis Gmelin, 1791, Patella... ee Calyptraeidae sinica Gmelin, 1791, Patella ........0000..ccececeeece neces Umbraculidae sinuosa Brocchi, 1814, Patella... Capulidae solandri Colenso, 1844, Patella .............0cccceececeeeeeeeee Acmaeidae spectrum Reeve, 1855, Patella ............c cc ceeceeeeeceeees Acmaeidae spinosa Gmelin, 1791, Patella .............. cece eeeeeeeeeee Fissurellidae squamulata Renier, 1804, Patella ........... . Calyptraeidae stella Lesson, 1831, Patella .......00.000ccccccceccceeeeeeeees Acmaeidae stellaris Reeve, 1855, Patella .......00...cccccccececcceeeeeee ees Acmaeidae stipulata Reeve, 1855, Patella ...............cccccccseeeeeeee Acmaeidae stowae Verco, 1906, Nacella ..........cc cece ..... Aemaeidae striata Reeve, 1855, Patella 0.0... eters Acmaeidae sturnus Hombron & Jacquinot, 1841, Patella .......... Acmaeidae subspiralis Carpenter, 1864, Nacella ..........0.00. Siphonariidae sulcata Borson, 1520, Patella ..... . Hipponicidae tectum Dillwyn, 1817, Patella tectumchinensis R6ding, 1798, Patella . Calyptraeidae . Calyptraeidae tenera C. B. Adams, 1845, Patella ......00000000000.... Acmaeidae tenuicostata Michelin, 1838, Patella ........000000000.0.... Acmaeidae tessellata O. F. Miiller, 1779, Patella .........0000000000... Acmaeidae testudinalis O. F. Miiller, 1776, Patella .......00.00.. Acmaeidae textilis Gould, 1846, Patella (Lottia) ........0000..000000. Acmaeidae tranquebarica Gmelin, 1791, Patella ........0.0.. Acmaeidae triangularis Carpenter, 1866, Nacella paleacea ....... Acmaeidae [01-562] November 27, 1973 tricarinata Linnaeus, 1767, Patella .........00...0.0.. Hipponicidae tricostata Gmelin, 1791, Patella oo... Hipponicidae trochiformis Gmelin, 1791, Patella ..................... Calyptraeidae trochoides Dillwyn, 1817, Patella .........00...0... Calyptraeidae tuberculifera Lamarck, 1819, Patella ................ ? Siphonariidae turcica Réding, 1798, Patella... Calyptraeidae umbellata Gmelin, 1791, Patella ...........00........... Umbraculidae umbellata della Chiaje, 1830, Patella ................. Umbraculidae umbonata Reeve, 1855, Patella .......00.....00...c00cece cece Acmaeidae uncinata Reeve, 1855, Patella ....................cccccecceeeees Acmaeidae undulata Réding, 1798, Patella oo... ee Caly ptraeidae ungarica Linnaeus, 1758, Patella .......0........ceeee Capulidae unguis Linnaeus, 1758, Patella ........0.0.....000....000eces. Fissurellidae unguis J. Sowerby, 1816, Patella... eee Capulidae unguisalmae Lesson, 1831, Patella .....00..000...0...000...08. Acmaeidae verriculata Reeve, 1855, Patella ....000....00000.cccecceeeeeee Acmaeidae vespertina Reeve, 1855, Patella ..........cceeeeeceeees Acmaeidae victoriae Gatliff & Gabriel, 1922, Patella ................ Acmaeidae victoriana Singleton, 1937, Patella ....0...00...cc eee. Acmaeidae virginea O. F. Muller, 1776, Patella .....00....000000...... Acmaeidae viridula Lamarck, 1819, Patella ..........00.......cc0cceees Acmaeidae zebrina Lesson, 1831, Patella o.....0.0...000cccccccceeeeeeeeee Acmaeidae Notes relevant to the above non-patellid species Patella hochstetteri K. Martin, 1879 Remarks—This species, from the Miocene of Java, is a Hemitoma, family Fissurellidae; in fact its author likened it to Hemitoma notata (Lin- naeus, 1755). Synonymy— 1879 Patella hochstetteri kK. Martin, Die tert. auf Java, Leiden, p. 86, pl. 12, fig. 10. Patella luchuana Pisbry, 1901 Remarks—Habe determined that the radula and gill structure of this Ryukyu Islands limpet prove it to belong to the Acmaeidae. Synonymy— 1901 Patella luchuana Pilsbry, Proce. Acad. Nat. Sci. Phila., vol. 53, p. 202. 1957 Collisella luchuana Pilsbry, Habe, Proc. Malac. Soc. Lond., vol. 32, p. 207. Patella nigrosulcata Reeve, 1855 Remarks—This shell, described from unknown locality, has since been identified as a Western Australian Patelloida, family Acmaeidae, and thus has nothing to do with Patella (Scutellastra) stel- laeformis, where it was assigned as a variety by Pilsbry, 1891. The species is usually found at- tached to the backs of large Patella (Scutellastra) laticostata Blainville. INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae 87 Synonymy— 1855 Patella nigro-sulcata| Reeve, Conch. Iconica. pl. 30, figs. 84 a, b. IS91 Patella (Scutellastra) stellaeformis var. nigrosulcata Reeve, Pilsbry, Man. of Conch., vol. 13. p. 100, pl. 66, figs. 66, 67. 1955 Patelloida nigrosulcata Reeve. Macpherson, Proc. Royal Society of Victoria, vol. 67 (2), p. 241. Patella opea Reeve, 1854 Remarks—Dr. Myra Keen has shown that the originally cited locality for this species, “Sand- wich Islands” - Hawaii, is incorrect, and that the species is a synonym of the West American Acmaea fascicularis Menke, 1851. Synonymy— 1854 Patella opea Reeve, Conch. Iconica, pl. 29, figs. 79 a, b. 1958 Acmaea fascicularis Menke, Keen, Sea Shells of Tropical West America, p. 244. Patella pallida Gould, 1859 Remarks—This Recent Japanese species is now recognised as belonging to the Acmaeidae. Synonymy— 1859 Patella pallida Gould, Proc. Boston Soc. Nat. Hist., vol. Ge 162: 1952 Tectura pallida Gould, Kuroda & Habe, Check List Rec. Mar. Moll. Japan, p. 89. Patella profunda Deshayes, 1863 Remarks—This Recent Reunion Island species is now known to belong to the Acmaeidae. Synonymy— 1863 Patella profunda Deshayes, Moll. Réunion, p. 44, pl. 6, figs. 15, 16. 1942 Patelloida profunda Deshayes, Tomlin & Stephenson, Proc. Malac. Soc., London, vol. 25, p. 6. Helcioniscus profundus var. mauritiana Pilsbry, 1891 Remarks—This Recent species and its variety from the island of Mauritius belong to the Ac- maeidae. Synonymy— IS9L Helcioniscus profundus var. mauritiana Pilsbry, 1891, Manual of Conchology, vol. 13, p. 150, pl. 65, figs 97-99. [01-563] SS Patellidae A. W. B. Powell List of Recognized Taxa SUBFAMILY Patellinae Genus Patella Linnaeus, 1758 Subgenus Patella Linnaeus, 1758 vulgata Linnaeus, 1758. Type. Europe aspera Roding, 1798. Europe depressa Pennant, 1777. Europe ferruginea Gmelin, 1791. Mediterranean baudonii Drouet, 1858. Azores rustica Linnaeus, 1758. S. Europe-Mediter- ranean piperata Gould, 1846. Madeira and Cape Verde Ids. caerulea Linnaeus, 1758. Mediterranean, Por- tugal, Azores and Canary Ids. moreleti Drouet, 1858. Azores lowei dOrbigny, 1839. Canary Ids. gomesti Drouet, 1858. Azores Subgenus (not known) candei d'Orbigny, 1839. Canary Ids. citrullus Gould, 1846. Madeira concolor Krauss, 1848. Natal and eastern South Africa depsta Reeve, 1855. St. Paul and Amsterdam Ids. rangiana Rochebrune, 1882. Cape Verde Ids. tkaffraria Rennie, 1930. Cretaceous, South Africa Subgenus Patellona Thiele, in Troschel & Thiele, 1891 granatina Linnaeus, 1758. Type. South Africa oculus Born, 1778. South Africa adansonii Dunker, 1853. West Africa canescens Gmelin, 1791. St. Helena lugubris Gmelin, 1791. West Africa and Cape Verde Ids. plumbea Lamarck, 1819. West Africa safiana Lamarck, 1819. Algeria to West Africa Subgenus Patellidea Thiele, in Troschel & Thiele, 1891 granularis Linnaeus, 1758. Type. South Africa Subgenus Cymbula H. & A. Adams, 1854 compressa Linnaeus, 1758. Type. South Africa miniata Born, 1778. South Africa Subgenus Olana H. & A. Adams, 1854 cochlear Born, 1778. Type. South Africa Subgenus Scutellastra H. & A. Adams, 1854 argenvillei Krauss, 1848. South Africa barbara Linnaeus, 1758. Type. South Africa longicosta Lamarck, 1819. South Africa tabularis Krauss, 1848. South Africa exusta Keeve, 1854. Mauritius subsp. pica Reeve, 1854. Mauritius to Sey- chelles flexuosa Quoy & Gaimard, 1834. Indo-Pacific, Andamans to Tuamotus subsp. optima Pilsbry, 1927. Japan kermadecensis Pilsbry, 1894. Kermadec Ids. taurorae Fleming, 1973. Middle Oligocene, New Zealand tucopiana (Powell, 1925). Tikopia, Melanesia laticostata Blainville, 1825. south West Australia peronii Blainville, 1825. southern Australia chapmani Tenison Woods, 1875. South Austra- lia to New South Wales thamiltonensis (Chapman Lower Pliocene, Australia tcooperi (Powell, 1938). Lower Miocene, New Zealand Subgenus Ancistromesus Dall, 1871 mexicana Broderip & Sowerby, 1829. Type. West Mexico t fuenzalidai Herm, 1969. Pliocene, Chile & Gabriel, 1923. Genus Helcion Montfort, 1810 Subgenus Helcion Montfort, 1810 pectunculus (Gmelin, 1791). Type. South Africa Subgenus Ansates Sowerby, 1839 pellucidus (Linnaeus, 1758). Type. Western Europe P tella (Bergh, 1871). Sargasso Sea Subgenus Patinastra Thiele, in Troschel & Thiele, 1891 pruinosus (Krauss, 1848). Type. South Africa dunkeri (Krauss, 1848). South Africa SUBFAMILY Nacellinae Genus Cellana H. Adams, 1869 eucosmia (Pilsbry, 1891). Red Sea radiata (Born, 1778). India to Philippines subsp. capensis (Gmelin, 1791). Natal to Zan- zibar subsp. enneagona (Reeve, 1854). Madagascar to Japan subsp. orientalis (Pilsbry, 1891). Indonesia; Japan; Marquesas tdeformis (K. Martin, 1883). Miocene, Java karachiensis (Winckworth, 1930). Gulf of Oman to Karachi livescens (Reeve, 1855). Type. Mauritius pricei Powell, new species. Samoa and New Hebrides garconi (Deshayes, 1863). Reunion and Mada- gascar [01-564] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae 89 Plate 65. Patella of the subgenus Scutellastra Figs. 1-3. Patella) (Scutellastra) barbara Linnaeus, 1758. Figs. 1, 3. Buluga Bay, East London, South Africa. Fig. 2. Port Alfred, South Africa. Fig. 4. Patella (Scutellastra) argenvillei: Krauss, 1548. Sea Point, South Africa. Figs. 5-7. Patella (Scutellastra) longicosta Lamarck, 1519. Kommetje, Cape Peninsula, South Africa. Figs. $-9. Patella (Scutellastra) flexuosa Quoy & Gaimard, 1834. Fig. 8. Paea, Tahiti. Fig. 9. Wake Island. Fig. 10. Patella (Scutellastra) flexuosa subspecies optima Pilsbry, 1927. Waki, Satsuma, Japan; young example. Figs. 11-13. Patella (Scutellastra) peronii Blainville, 1825. Fig. 11. Swansea, Tasmania. Figs. 12-13. Shellharbour, New South Wales, Australia. [01-565] 90 - Patellidae (Cellana cont'd) testudinaria (Linnaeus, 1758). Andaman Ids. to New Caledonia vitiensis Powell, new name. Fiji grata (Gould, 1859). Japan and Korea mazatlandica (Sowerby, 1839). Japan and Ry- ukyu Ids. nigrolineata (Reeve, 1854). Japan toreuma (Reeve, 1855). Japan to Philippines exarata (Reeve, 1854). Hawaiian Ids. talcosa (Gould, 1846). Hawaiian Ids. tahitensis (Pease, 1868). Tahiti and Pitcairn ardosiaea (Hombron & Jacquinot, 1841). Juan Fernandez Id. conciliata Iredale, 1940. Queensland turbator Iredale, 1940. South Queensland tramoserica (Holten, 1802). South Queensland to South Australia solida (Blainville, 1825). Tasmania to South Australia tcarpentariana Skwarko, 1966. Lower Creta- ceous, North Australia tcudmorei Chapman & Gabriel, 1923. Lower Mi- ocene, Victoria thentyi Chapman & Gabriel, 1923. Lower Plio- cene, Victoria analogia Iredale, 1940. Lord Howe Id. howensis Iredale, 1940. Lord Howe Id. craticulata (Suter, 1905). Kermadec Ids. denticulata (Martyn, 1784). New Zealand flava (Hutton, 1873). New Zealand ornata (Dillwyn, 1817). New Zealand radians (Gmelin, 1791). New Zealand stellifera (Gmelin, 1791). New Zealand strigilis (Hombron & Jacquinot, 1841). Auck- land and Campbell Ids. subsp. bollonsi Powell, 1955. Antipodes Ids. subsp. chathamensis (Pilsbry, 1891). Chatham Ids. subsp. flemingi Powell, 1955. Snares Ids. subsp. oliveri Powell, 1955. Bounty Ids. subsp. redimiculum (Reeve, 1854). Southern New Zealand + thomsoni Powell & Bartrum, 1929. Lower Mio- cene, New Zealand tcophina Powell, new species. Lower Miocene, New Zealand taberna Powell, new species. Lower Miocene, New Zealand 3 Genus Nacella Schumacher, 1817 Subgenus Nacella Schumacher, 1817 mytilina (Helbling, 1779). Type. Southern Chile to Kerguelen Id. A. W. B. Powell kerguelenensis (E. A. Smith, 1877). Kerguelen and Heard Ids. Subgenus Patinigera Dall, 1905 clypeater (Lesson, 1831). Chile concinna (Strebel, 1908). South Georgia to Antarctica deaurata (Gmelin, 1791). Patagonia, Falklands, Tierra del Fuego subsp. delicatissima (Strebel, 1907). Magellan and Falklands delesserti (Philippi, 1849). Marion Id. edgari (Powell, 1957). Kerguelen Id. flammea (Gmelin, 1791). Strait of Magellan fuegiensis (Reeve, 1855). Magellan, Falklands, South Georgia magellanica (Gmelin, 1791). Type. Magellan to Falklands subsp. venosa (Reeve, 1854). Chiloe Island, Chile macquariensis Finlay, 1927. Heard Ids. terroris (Filhol, 1880). Campbell Id. Macquarie and Acknowledgements The writer is greatly indebted to Mr. W. B. Dixon Stroud for his continued generous mon- etary support of this and other projects, intended for publication, or already published, in “Indo- Pacific Mollusca.” To the following people who have assisted with information, photographs, the loan of types, and other material, the writer gratefully acknowledges the help afforded by—Dr. R. T. Abbott, Delaware Museum of Natural History; Professor Dr. W. Adam, Institut Royal des Sciences Naturelles de Belgique; Mr. W. O. Cernohorsky, Auckland In- stitute and Museum; Dr. F. M. Climo, Dominion Museum, Wellington; Mr. N. W. Gardner, Auck- land; Mrs. J. Kerslake, Sydney; Dr. Y. Kondo, B. P. Bishop Museum, Honolulu; Mr. I. G. Mar- row, Melbourne; Dr. D. F. McMichael, formerly Australian Museum, Sydney; Mrs. V. Orr Maes, Academy of Natural Sciences of Philadelphia; Dr. J. F. Peake, British Museum (Natural His- tory); Dr. W. F. Ponder, Australian Museum, Sydney; Mr. L. Price, Kaitaia, New Zealand; Dr. H. A. Rehder, United States National Museum; Dr. J. D. Taylor, British Museum (Natural His- tory); and the late Mr. D. Thaanum, Honolulu. To Mrs. Nancy Prior of Cape Town, the writer is especially indebted for the fine examples of South African limpets provided for the colour plates. [01-566] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae 91 Selected Bibliography Anton, H. E. 1839. Verzeichniss der Conchylien welche sich in der Sammlung von Hermann Eduard Anton betinden. Halle, pp. 1-110. Blainville, H. M. D. 1825. Dictionaire des Sciences Naturelles, vol. 38. Born, I. von. 1778. Index rerum naturalium Musei Caesarei Vindobonensis, Testacea. Vienna, pp. 1-458. Brian, M. V. and Owen, G. 1952. The Relation of the radula fraction to the environment in Patella. Jour. of Anim. Ecol- ogy, vol. 21 (2), pp. 241-249. 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B. and Jacquinot, C. H., 1841. Suite de la de- scription de quelques Mollusques, provenant de la cam- pagne de l’Astrolabe et de la Zélée. Annales des Sciences Naturelles, Zool. & Paleont. ser. 2, vol. 16, pp. 190-192. Iredale, T. 1940. Marine Mollusca from Lord Howe Island, Norfolk Island, Australia, and New Caledonia. Australian Zoologist, vol. 9, pt. 4, pp. 432-433 (Patellidae). Jay, J. C. 1839. Catalogue of Recent Shells in the Cabinet of J. C. Jay, 3rd. ed., pp. 1-125, New York. Kobelt, W. 1886-1887. Prodromus Faunae Molluscorum Testaceorum maria europaea inhabitantium. Niirnberg. Fasc. 1, pp. 1-128 (1886); Fase. 2, pp. 129-256 (1887); Fasc. 3, pp. 257-384 (1887); Fasc. 4, pp. 385-550 (1887). Koch, H. J. 1949. A Review of the South African Representa- tives of the Genus Patella Linnaeus. Annals of the Natal Museum, vol. 11, pt. 3, pp. 487-517. Knight, J. Brooks, et al. 1960. Moore’s Treatise on Paleontol- ogy. Part 1, Mollusca 1, pp. 231-237 (Patellacea). Krauss, F. 1848. Die Sudafrikanischen Mollusken, Stuttgart, pp. 1-140. Lesson, R. P. 1831. Voyage Autour du Monde, éxecuté par ordre du Roi, sur la corvette de La Majesté, La Coquille, pendant les années 1822, 1823, 1824 and 1825. Zoologie. Paris, vol. 2 (1), pp. 239-471. Lightfoot, J. 1786. A Catalogue of the Portland Museum, lately the Property of the Duchess of Portland, London, pp. 1-194. Linnaeus, C. 1758. Systema naturae per regna tria naturae. 10th ed. Stockholm, pp. 1-824. Lowenstam, H. A. 1962. Geothite in Radular teeth of Recent Marine Gastropods. Science, vol. 137, No. 3526, pp. 279- 280. Lowenstam, H. A. 1962. Magnetite in Denticle Capping in Recent Chitons (Polyplacophora). Geol. Soc. America, Bull. 73, pp. 435-435. Macpherson, J. Hope. 1955. Preliminary Revision of the Fam- ilies Patellidae and Acmaeidae in Australia. Proc. Royal Soc. Victoria, vol. 67, pt. 2, pp. 229-256. Martyn, T. 1784-1786. The Universal Conchologist. London, vols. | and 2, pls. 1-80 (1754); vols. 3 and 4, pls. 51-160 (1786). Mermod, G. 1950. Les Types de la Collection Lamarck au Museum de Geneve. Mollusques vivants, 1. Revue Suisse de Zoologie, vol. 57, no. 34, pp. 689-701 (Patella). Michelotti, G. 1847. Description des Fossiles des Terrains Miocenes de I'Italie septentrionale. Holl. Maat. Wetensch. Nat. Verh., vols., 2, 3 (2). Haarlem. Nordsieck, F. 1968. Die europaischen Meeres-Gehiusesch- necken (Prosobranchia) Vom Eismeer Kapverden und Mittelmeer. Stuttgart. pp. 13-15 (Patellidae). Oliver, W. R. B. 1915. The Mollusca of the Kermadec Islands. Trans. New Zealand Inst., vol. 47, pp. 310-314 (Patellidae). Orton, J. H. 1928. Observations on Patella vulgata, pt. 1: Sex- phenomena, breeding and _ shell-growth. Jour. Mar. Biol. Assoc. U. K., vol. 15, pp. 851-862. Orton, J. H. 1928. Observations on Patella vulgata, pt. 2: Rate of growth of shell. Jour. Mar. Biol. Assoc. U. K., vol. 15, pp. 863-874. Orton, J. H. 1929. Observations on Patella vulgata, pt. 3: Habitats and Habits. Jour. Mar. Biol. Assoc. U. K., vol. 16, pp. 277-288. Pallary, P. 1912. Exploration scientifique du Maroc. Mala- cologie. Institut scientifique chérifien, Fasc. 12, pp. 1-107. Payraudeau, B. C. 1826. Catalogue descriptif et methodique des Annelides et des Mollusques de I'Ile de Corse, Paris, 8 pls. [01-571] 92 Patellidae A. W. B. Powell Perry, G. 1811. Conchology, or the natural history of shells. London. pl. 43 (Patella). Pilsbry, H. A. 1891. Acmaeidae, Lepetidae, Patellidae and Titiscaniidae. Manual of Conchology, vol. 13, pp. 1-195. Powell, A. W. B. 1951. Antarctic and Subantarctic Mollusca: Pelecypoda and Gastropoda. Discovery Rep. vol. 26, pp. 80-84 (Patellidae). Powell, A. W. B. 1955. Mollusca of the Southern Islands of New Zealand. Cape Exped. Ser., Dept. Sci. Indust. Res., Bull. 15, pp. 65-74 (Patellidae). Powell, A. W. B. 1957. Mollusca of Kerguelen and Macquarie Islands. British, Australian, New Zealand Antarctic Research Expedition, 1929-1931, ser. B, vol. 6, pt. 7, pp. 126-128 (Patellidae). Reeve, L. A. 1854-1855. Monograph of the genus Patella. Conchologia Iconica, pls. 1-42, text and index. Pls. 1-24, sp. 1-64 dated 1854; pls. 25-42, sp. 65-144 dated 1855. Réding, P. F. 1798. Museum Boltenianum, pt. 2, Hamburg, pp. 1-12 (Patella). Schaffer, F. X. 1912. Das Miocan von Eggenburg. Abh. geol. Reichs. Anst., vol. 22 (2), pp. 129-193. Schuster, E. 1913. Anatomy of Helcioniscus ardosiaeus H. & J. Zool. Jahrb., Jena, Suppl. 13, Fauna Chilensis, vol. 4, pp. 281-284. Smith, F. G. W. 1935. The Development of Patella. Phil. Trans. series B., vol. 225, pp. 95-125. Stephenson, T. A. 1948. The Constitution of the Intertidal Fauna and Flora of South Africa. Part .3. Annals of the Natal Museum, vol. 11, pt. 2, pp. 207-324. Strebel, H. 1907. Beitrage zur kenntnis der Molluskenfauna der Magalhaen-Provinz, No. 5, pp. 110-155; pls. 3-7 (Patel- lidae). Thiele, J. in Troschel & Thiele, 1891. Das Gebiss der Schnecken, vol. 2 (7), pp. 309-334; pl. 28 (Patellidae). Thiem, H. 1917. Beitrage zur Anatomie und Phylogenie der Docoglossen. 1. Zur Anatomie von Helcioniscus ardosiaeus Hombron et Jacquinot unter Bezugnahme auf die Bear- beitung von Erich Schuster in den Zoolog. Jahrb., Supple- ment 13, vol. 4, 1913. Jena Z. Naturw. vol. 54, pp. 333-404. Thomson, J. A. 1919. Polymorphism in the Common New Zealand Limpet, Cellana radians (Gmelin). N. Z. Journ. Sci. & Tech., vol. 2, pp. 264-267. Tomlin, J. R. le B. and Stephenson, T. A. 1942. South African Patellidae. Proc. Malac. Soc. London, vol. 25, pt. 1, pp. 4-9. Turton, W. H. 1932. The Marine Shells of Port Altred S. At- rica. Oxford Univ. Press, pp. 160-171 (Patellidae). Wood, W. 1828. Supplement to the Index Testaceologicus; or a catalogue of shells, British and Foreign, London, pp. 1-59. [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence. | (01-5 -~l bo ] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae 93 Plate 66. Patella of the subgenus Scutellastra Fig. 1. Patella (Scutellastra) kermadecensis Pilsbry, 1894. Figs. 3-4. Patella (Scutellastra) laticostata Blainville, 1825. Raoul Island, Kermadec Islands. Largest living species of the Albany, Western Australia. subgenus. Fig. 5. Patella (Scutellastra) tabularis Krauss, 1848. Cape Fig. 2. Patella (Scutellastra) flexuosa subspecies optima Point, South Africa. Pilsbry, 1927. Yakushima, Japan. [01-577] 94 Patella Family Patellidae Rafinesque 1815 The Patellidae or family of true limpets have simple, oval to rounded, conical or cap-shaped shells, without a perforation, marginal notch or internal septum. They are characteristic of the intertidal zone and seldom extend much below low-tide mark. A detailed account of the animal, its habits, functions, distribution and geological range, is given in the introductory section of this work. Subfamily Patellinae Rafinesque, 1815 The subfamily Patellinae includes the genera Patella and Helcion, as well as several subgenera of each. The radula comprises four identical central teeth, often with the addition of a median central that may vary from vestigial to fully developed. The lateral is large and pluricuspid, and_ is flanked by three weak, slender, apparently func- tionless marginals. The radular ribbon is relatively short, straight, and folded back upon itself at the nascent end. The gill cordon is continuous in all members, except in typical Helcion, which has the cordon interrupted by the head, understandable in that instance, since the sole species, pectunculus, has the anterior end reduced almost to nothing. The shell in Patella is usually rather solid, por- cellanous within, and seldom iridescent. On the other hand, Helcion (Patinastra) is semitrans- parent, and Cellana-like, except for the dentition which closely resembles that of Patella. The typical genus, Patella, is distributed along most of the eastern coast of Europe, from the Lofoten Islands, and including Britain, to Spain, the Mediterranean, west coast of Africa and off- shore islands; also there is one species in Natal, and another, somewhat atypical, at the South Indian Ocean Islands of St. Paul and Amsterdam. The subgenus Patellona is predominantly West African, but extends to South Africa; the sub- genera Cymbula and Olana are exclusively South African; the subgenus Scutellastra is South A fri- can as well, but also has a very extensive Indo- Pacific range, and the subgenus Ancistromesus, A. W. B. Powell Patellidae largest of all limpets, belongs exclusively to the west coast of Central America. Numerous species, attributed to Patella, Hell- cion and Nacella, have been described from European Cretaceous and Tertiary horizons, but their true identity, of necessity based upon shell characters alone, is uncertain. Genus Patella Linnaeus, 1758 Type: Patella vulgata Linnaeus, 1758 Shell ovate, conical or cap-shaped, with the apex subcentral, usually solid, and of medium size to very large. Sculpture consisting of radial ridges of varying strength, mostly crossed by concentric growth lines. Interior of shell varying from sub- translucent, polished and _ iridescent, to opaque porcelanous. Colour pattern external, usually in the form of radials associated with the ribbing, and showing through to the interior in subtrans- lucent shells but confined to the marginal border in those with a thick porcelanous internal layer. The gill cordon is complete, and the radula relatively short and folded back upon itself at the nascent end. The radula formula is— 341444143 oF ot 14 (24 Le 2) 1+ sor SPO De 3 The variations of the above formulae occur in the central teeth, which may consist of 4 iden- tical centrals in a horizontal row, as in Patella vulgata, or in others when an incipient median central is added, or, again, in certain species of the subgenus Scutellastra when the median cen- tral attains the size of the other centrals, thus making 5 identical centrals. The lateral is almost invariably pluricuspid, and the 3 marginals are narrow, with very weak cusps at most, and they are apparently functionless. The range of the genus is wide-spread in warm and temperate seas, but is absent from certain regions, notably both coasts of North America, the Caribbean and South America. A number of fossil species attributed to Patella has been described, ranging from the upper Cre- taceous onward, but most of these are difficult to assign generically or even to family since we lack knowledge of the soft parts. The genus Patella is here divided into several subgenera that are each more or less restricted to definite geographical areas. Their synonymy is recorded under the relevant subgenera. [01-578] November 27, 1973 Subgenus Patella Linnaeus, 1758 Type: Patella vulgata Linnaeus, 1758 Shell of small to moderate size, the inner layer subtranslucent and more or less iridescent, often with the external colour pattern showing through the glaze. Gill cordon complete and radula with 4 identical central teeth, arranged in a horizontal row, and occasionally with an incipient median central, represented by a narrow functionless plate. Distribution, the western coastline of Europe, from the Lofoten Islands, and including Britain, to the Mediterranean, down to Madeira and the Canary Islands, and appearing again along the coast of Natal. Synonymy— 1758 Patella Linnaeus, Syst. Nat., ed. 10, p. 780. Type, by subsequent designation, Fleming, 1818: Patella’ vul- gata Linnaeus, 1755S. 1810 Patellus Montfort, Conchyliologie Systématique, vol. 2, p. 67. Type, by original designation: Patellus roseus Montfort, 1810. 1884 Patellopsis Thiele in’ Troschel, Das Gebiss der Schnecken, vol. 2, p. 324, based upon the radula (pl. 28, fig. 22) of an unnamed South African Patella, pos- sibly variabilis Krauss, 1848. 1884 Patellastra Monterosato, Natural. Sicil., vol. 3, p. 103. Type, by monotypy: Patella lusitanica Gmelin, 1791. 1912 Costatopatella Pallary, Mem. Inst. Egypte, vol. 7 (3), p. 148. 1920 Granopatella Pallary, Arch. Sci. Prot. Franc. Expl. Sci. Maroc., fasc. 2, p. 72. 1920 Laevipatella Pallary, Arch. Sci. Prot. Franc. Expl. Sci. Maroc., fase. 2, p. 72. Patella vulgata Linnaeus, 1758 (Pl. 63, figs. 1-3; pl. 68, figs. 1, 2; pl. 61, fig. 1) Range—Western Europe, Lofoten Islands to Spain and the British Isles. Remarks—This is the common European edible limpet. It is moderately large, solid, oval and conical, radially ribbed, and usually whitish or yellowish, often radially lined or streaked in brown. Description—Shell moderately large, up to 60 mm. (2% inches) in length, solid, oval, conical, with the apex a little in front of the middle, and sculptured with radiating ribs and interstitial lirae. Colour varying from whitish to yellowish, sometimes radially lined or streaked with dark- brown. Interior weakly iridescent, the spatula grayish to leaden colour or clouded with whitish INDO-PACIFIC MOLLUSCA, vol. 1, no. 15 Patella 95 callus, often with the shell margin dark-lined by the external pattern showing through. Radula—Formula 3+1+4+1+3. The four central teeth are of approximately equal size, and are arranged in a straight horizontal line, without a median vestigial central. Measurements (mm.)— length width height 60.5 53.0 32.0 Caldy Island, S$. Wales 43.0 O75 21.0 Isle of Man Synonymy— 1758 Patella vulgata Linnaeus, Syst. Nat., ed. 10, p. 752. ? 1798 Patella conus Réding, Mus. Bolten., pt. 2, p. §. 1811 Patella radiata Perry, Conch., London, pl. 43, fig. 1 (non Born, 1778). 1839 Patella conica Anton, Verzeichniss, p. 26 (non Blain- ville, 1825). 1844 Patella vulgata var. conica Brown, Illust. Conch., ed. 2, p. 65. 1844 Patella vulgata var. communis Brown, Illust. Conch., ed. 2, p. 63. 1854 Patella vulgata Linn., Reeve, Conch. Iconica, vol. 8, pl. 18, figs. 42 a-c. (Dec.). 1857 Patella vulgata var. intermedia Knapp (in Murray), Ann. Mag. Nat. Hist., 19, p. 211. 1865 Patella vulgata var. elevata Jeffreys, Brit. Conch., vol. 3, p. 237. 1865 Patella vulgata var. picta Jeffreys, Brit. Conch., vol. 3, p. 237. 1887 Patella vulgata var. secernenda Dautzenberg, Excur. mal. St.-Lunaire, p. 13. 1891 Patella vulgata Linn., Pilsbry, Man. Conch., vol. 13, p. 52, pl. 10, figs. 1-6. 1906 Patella vulgata var. aurea Martel in Dautzenberg & Durouchoux, Suppl. Faun. malac. St.-Malo, p. 11. 1906 Patella vulgata var. major Dautzenberg & Durouchoux, Suppl. Faun. malac. St.-Malo, p. 11. Patella ferruginea Gmelin, 1791 (PI. 69, figs. 1-3) Range—Mediterranean, from the Aegean to Spain and North Africa. Remarks—This species is easily recognised by its thick shell, strong radial ribs, deeply corrugated margin and ashen colour. Description—Shell moderately large, up to 62 mm. (2-7/16 inches) in length, very solid, ovate, conical, with the apex subcentral, coarsely sculp- tured with numerous strong radial ribs, that are rendered scabrous by concentric growth lines, and also strongly corrugate the margin. Colour, extern- ally dull ashen, more or less stained with pale brown; internally, bluish white, corrugated margin bordered in dark-brown, almost black, and the spatula clouded with whitish callus. Radula—Formula 3 + 1 + (2+1+2) + 1+ 3. Radula very similar to that of caerulea, except that the [01-579] 96‘ Patella small slender median central is a definite tooth bearing a small cusp. Measurements (mm. )— length width height 60.0 51.5 24.0 Corsica 59.0 47.0 21.0 Corsica Synonymy— 1791 Patella ferruginea Gmelin, Syst. Nat., ed. 13, p. 3706; based upon Martini-Chemnitz, Conch. Cab., vol. 1, pl. 8, tig. 66. 1819 Patella luteola Lamarck, Anim. sans vert., vol. 6 (1), p. 327. 1819 Patella pyramidata Lamarck, Anim. sans vert., vol. 6 (1), p. 327. 1826 Patella rouxii Payraudeau, Cat. Moll. Corse, p. 90. 1826 Patella lamarckii Payraudeau, Cat. Moll. Corse, p. 90. 1854 Patella costoso-plicata Reeve, Conch. Iconica, vol. 8, pl. 8, figs. 14 a, b. 1884 Patella ferruginea var. ficarazzensis de Gregorio, Bull. Soc. Mal. Ital., vol. 10, pp. 120-124. 1884 Patella ferruginea var. imperatoria de Gregorio, Bull. Soc. Mal. Ital., vol. 10, pp. 120-124. 1884 Patella ferruginea var. percostata de Gregorio, Bull. Soc. Mal. Ital., vol. 10, pp. 120-124. 1884 Patella ferruginea var. sitta de Gregorio, Bull. Soc. Mal. Ital., vol. 10, pp. 120-124. 1891 Patella ferruginea Gmelin, Pilsbry, Man. Conch., vol. 13, p. 81, pl. 53, figs. 1-3; pl. 17, figs. 23, 24. 1950 Patella luteola Lamarck, Mermod, Rev. Suisse Zool., vol. 57, no. 34, p. 692, fig. 3 (type). 1968 Patella ferruginea Gmelin, Nordsieck, Eur. Meeres- Gehauseschn. Stuttgart, p. 15. Records—CORSICA; near Bonifacio (AWBP. coll. 28388). SPANISH MOROCCO; Melilla (AWBP. coll.); Chafarinas Is- lands (Zafarines), 35° 10’ N., 2° 25’ E. (AWBP. coll.). Types—The type of luteola is in the Museum d'Histoire Naturelle de Geneve. Patella baudonii Drouet, 1858 (Pl. 75, figs. 1, 2) Range—Azores, Santa Maria and Pico. Remarks—This species, which the writer has not seen, seems to be closely allied to, if not identical with, Patella ferruginea Gmelin, 1791. Pilsbry’s translation of the original description follows, and the illustrations are from Drouet’s original figures. Description—‘Shell large, subelevated, coarsely ribbed, plicate, solid, thick; outside greyish-green, inside white; vertex subacute, submedian; aper- ture oval, a little crenated.” Measurements (mm. )— length width height 60.0 50.0 25.0 (Drouet) A. W. B. Powell Patellidae Synonymy— 1858 Patella baudonii Drouet, Moll. Mar. Agores, p. 41, pl. 2, figs. 8, 9. 1891 Patella baudonii: Pilsbry, Man. Conch., vol. 13, p. 86, pl. 54, figs. 15, 16. Patella rustica Linnaeus, 1758 (Pl. 69, figs. 4, 5) Range—Atlantic coast of south west France, Portugal, Spain, Mediterranean and Adriatic Seas. Remarks—This species, better known by the Gmelin name, lusitanica, is rather small, ovate- conical, and densely sculptured with fine granular radials. The external colour is greyish, or pale brownish, speckled with black, and internally it is broadly radially banded in dark puplish-brown. A nearly related species is the narrowly-ovate piperata from Madeira and the Cape Verde Islands. Description—Shell rather small, up to 35 mm. (1% inches) in length, solid, ovate, tall-conical, with the apex slightly anterior to the middle. Sculpture con- sisting of very numerous, closely spaced, narrow, somewhat uneven, granulose radial riblets. Colour, externally pale yellowish-brown to greyish, often with the rib-granules black, internally broadly rayed in dark-brown or blue-black on a greyish- silvery ground. Spatula white callused, often sur- rounded by a yellowish-brown stain. Radula—F ormula 3 + 1 + (2+1+2) + 1+ 3. Radula with or without a narrow median central tooth, remaining four centrals of uniform size, and ar- ranged in a horizontal line. Measurements (mm.)— length width height 35.0 28.5 17.0 Melilla, Morocco 28.0 24.0 12.55. Melilla, Morocco Synonymy— 1758 Patella rustica Linnaeus, Syst. Nat., ed. 10, p. 783. 1791 Patella lusitanica Gmelin, Syst. Nat., ed. 13, p. 3715. 1798 Patella squamata Roding, Mus. Bolten, pt. 2, p. 10. 1819 Patella punctata Lamarck, Anim. sans vert., vol. 6, p. 333. 1825 Patella subgranularis Blainville, Dict. Sci. Nat., vol. 38, p. 113. (fide Christiaens, 1968, p. 367). 1854 Patella nigro-punctata Reeve, Conch. Iconica, vol. 8, pl. 23, figs. 57 a-c (Dec.). 1883 Patella lusitanica var. minor Marion, Faune bass. med., p. 48. 1884 Patellastra lusitanica Gmel., Monterosato, Natural. Sicil., vol. 3, p. 103. 1891 Patellastra lusitanica Gmel., Thiele, in Troschel & Thiele, Das Gebiss der Schnecken, 2, pl. 28, fig. 12 (radula). 1891 Patella lusitanica Gmel., Pilsbry, Man. Conch., vol. 13, p. 87, pl. 11, figs. 15-19. 1912 Patella rustica Linné var. major Pallary, Explor. scient. Maroc., p. 72. [01-580] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 97 Plate 67. Indo-Pacitic Cellana Figs. 1, 2. Cellana testudinaria (Linnaeus, 1758). Vanualava, Banks Islands. Fig. 3. Cellana talcosa (Gould, 1846). Molokai, Hawaiian Islands. Figs. 4-6. Cellana exarata (Reeve, 1854). Molokai, Hawaiian Islands. Fig. 7. Cellana grata (Gould, 1859). Matsushima Island, Korea. Figs. 8,9. Cellana radiata (Born, 1778). Colombo, Ceylon. Figs. 10, 11. Cellana radiata subspecies capensis (Gmelin, 1791). Near Durban, Natal. Figs. 12, 13. Cellana radiata subspecies orientalis (Pilsbry, 1891). Fig. 12. Russell Islands, Solomon Islands (note the strong radial folds). Fig. 13. Tau Island, Samoa. Fig. 14. Cellana radiata subspecies enneagona (Reeve, 1854). Jolo, Philippine Islands. Figs. 15, 16. Cellana nigrolineata (Reeve, 1854). Fig. 15. Fukura, Awaji, Japan. Fig. 16. Chiringashima, Japan. Figs. 17, 18. Cellana mazatlandica (Sowerby, 1539). Bonin Islands. [01-581] 98 Subgenus Patella A. W. B. Powell Patellidae Plate 68. Figs. 1, 2. Patella vulgata Linnaeus, 1758. Fig. 1. Kimmeridge, England, 52 mm., AWBP coll. 11358. Fig. 2. Isle of Man, 38 mm., AWBP coll. 11359. Figs. 3, 4. Patella aspera Roding, 1798, Caldy Island, Wales, 47 mm., AWBP coll. 217. 1912 Patella rustica Linneé var. maroccana Pallary, Explor. scient. Maroc., p. 72. 1938 Patella lusitanica var. orientalis Pallary, Jour. Conchyl., vol. 82, p. 47. 1950 Patella punctata Lam., Mermod, Rev. Suisse Zool., vol. 57 (34), p. 695, fig. 7 (type). 1968 Patella lusitanica Gmelin, Christiaens, Bull. Mus. Nat. WHist. Nat., ser. 2, vol. 40 (2), pp. 366, 367. 1968 Patella (Patellastra) rustica L., Nordsieck, Die europ- Meeres Gehauseschnecken, Stuttgart, p. 15. Patella piperata Gould, 1846 (Pl. 71) Range—Madeira and Cape Verde Islands. Remarks—Shell very similar to that of rustica in sculpture and in coloration, but more elongate-ovate in its younger stages, and with the apex nearer to the anterior end. Radula—The radula differs from that of rustica in that the four central teeth are not in horizontal alignment, the middle pair being set lower than the outer pair (Christiaens, 1968, p. 370, fig. 2a). Plate 69. Figs. 1-3. Patella ferruginea Gmelin, 1791. Fig. 1. Chafarines Islands, Morocco, 55 mm., AWBP. coll. 1054. Fig. 2. Bonifacio, Corsica, 60 mm., AWBP coll. 28388. Fig. 3. Melilla, Morocco, 42 mm., AWBP coll. 30974. Figs. 4, 5. Patella rustica Linnaeus, 1758, Oran, Algeria, 25-26 mm., AWBP coll. 80. Measurements (mm.)— length width height 44.0 40.0 — Christiaens, 1968, p. 372 27.0 21.0 12.0 Madeira Synonymy— 1839 Patella guttata Orbigny, in Webb & Berthelot, Hist. Nat. Moll. Canaries, p. 98 (non Gmelin, 1791). 1846 Patella piperata Gould, Proc. Boston Soc. Nat. Hist., vol. 2, p. 150. 1846 Patella nigrosquamosa Dunker, Zeitschr. f. Malak., p. 25. 1866 Patella frauenfeldi Dunker, Verh. k. k. zool.-bot. Ges. Wien, vol. 16, p. 914. “Madras” in error for Madeira. 1867 Patella frauenfeldi Dunker, Frauenfeld, Reise Novara, Zool., vol. 2, pt. 3, Moll., p. 15, pl. 2, figs. 26 a, b. 1968 Patella piperata watsoni Christiaens, Bull. Mus. Nat. @Hist. Nat. ser. 2, vol. 40, no. 2, p. 371, text fig. 2 b; pl. 1, fig. b. 1968 Patella piperata nigro-radiata Christiaens, Bull. Mus. Nat. @Hist. Nat. ser. 2, vol. 40, no. 2, p. 371, text fig. 2 c; pl. 1, fig. c. 1968 Patella piperata alba Christiaens, Bull. Mus. Nat. d’Hist. Nat. ser. 2, vol. 40, no. 2, p. 371, pl. 1, fig. g (non P. alba Anton, 1539). [01-582] November 27, 1973 Patella caerulea Linnaeus, 1758 (Pls. 61, 63, 72, 74) Range—Mediterranean and Adriatic Seas, Por- tugal, Azores, Madeira and Canary Islands. Remarks—This species is variable in shape, col- our, and strength of the radial ribbing, but in gen- eral terms it is a depressed, thin, and spreading shell, with 6 or 7 distinct marginal angles, result- ant from 7 to 9 prominent radial folds. The typical form of the species from the Mediterranean and Adriatic Seas, has a colour range, varying from al- most white to buff or pale brownish, often radially banded with iridescent blue. Shells from the Azores and Madeira are larger, even more depressed than the typical species, have broader and more prominent radial folds, and are of much darker colour, being dark reddish brown externally, similarly coloured internally, but dif- fused with iridescent blue, and with a distinct- edged, white spatula. This latter form is crenata Gmelin, and when more material is studied, Gmelin’s name may prove to be usable to define a regional subspecies of caerulea, restricted to the Azores, Madeira and Canary Islands. Negating this possibility is the fact that shells from the ad- jacent mainland of Spanish Morocco have the dark colouring of crenata but a shape and sculpture similar to those in typical caerulea. Description—Shell of moderately large size, 40-71 mm. (1/2-2% inches) in length, thin, depressed, usu- ally distinctly 6 or 7 angled, resultant from 7 to 9 broadly rounded primary folds that project at the margin. Surface crowded with secondary radials of varying sizes, mostly imbricated by concentric growth lines. Colour whitish or buff externally, the interior silvery-white, radially lined or banded in blue, the spatula bluish or white-callused. Some examples have a pale yellowish interior without radial markings, and the form crenata is dark red- dish brown, internally diffused with iridescent blue, and with a clear-cut white spatula. Radula—Formula 3 + 1 + (2+X+2) + 1+ 3. The radula is of the same style as that of vulgata, except for a slight median gap between the pairs of four centrals, in which appears a narrow vestigial plate. The four functional centrals, as in vulgata, are in a straight horizontal row. INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 99 Measurements (mm.)—(all A. W. B. Powell collection.) length width height 51.0 46.0 12.0 Capri, Italy 36.0 30.0 10.5 Melilla, Morocco 71.0 64.0 14.0 Madeira (crenata form) Synonymy— 1758 Patella caerulea Linnaeus, Syst. Nat., ed. 10, p. 782. 1791 Patella crenata Gmelin, Syst. Nat., ed. 13, p. 3706. 1791 Patella margaritacea Gmelin, Syst. Nat., ed. 13, p. 3707. 1793 Patella tarentina von Salis, Reise ins. Koenig. Neapel, p. 359, pl. 6, fig. 2. 1798 Patella silicina Riding, Mus. Bolten., pt. 2, p. 9. 1819 Patella tarentina Lamarck, Anim. sans vert., vol. 6, p. 332. 1826 Patella bonnardii Payraudeau, Moll. de Corse, p. 89. 1836 Patella fragilis Philippi, Enum. Moll. Sicil., vol. 1, p. 110. 1838 Patella subplana Potiez & Michaud. Gal. Moll. Douai, vol. 1, p. 524. 1854 Patella scutellaris Lam., Reeve, Conch. Iconica, vol. 8, pl. 20, fig. 49. 1882 Patella stellata Bucquoy, Dautzenberg & Dollfus, Moll. mar. Roussillon. (non Helbling, 1779). 1882 Patella adspersa Bucquoy, Dautzenberg & Dollfus. Moll. mar. Roussillon. 1882 Patella caerulea var. cognata Bucquoy, Dautzenberg & Dollfus, Moll. mar. Roussilon, p. 471. 1882 Patella caerulea var. intermedia Bucquoy, Dautzenberg & Dollfus, Moll. mar. Roussilon, p. 471. 1891 Patella caerulea Linne~ Pilsbry, Man. Conch., vol. 13, p. 83, pl. 10, figs. 7-12. 1950. Patella tarentina Lam., Mermod, Rev. Suisse Zool., vol. 57 (34), p. 695 (text figs. of type series). Records—ITALY: Naples; Isle of Capri; Palermo, Sicily. MAL- TA. MOROCCO: Melilla. MADEIRA (crenata form). (All AWBP coll.) Patella moreleti Drouet, 1858 (Pl. 75, figs. 3, 4) Range—Fayal, Azores. Remarks—The writer has not seen this species which possibly may be only a form of Patella caerulea Linnaeus. Pilsbry’s translation of the orig- inal description follows, accompanied by copies of Drouet’s figures. Description—‘Shell subdepressed, very rugose, ribbed, the ribs scaly, scarcely solid; brownish- green outside; inside brownish or reddish, irides- cent, with a white spot at the summit. Apex acute. Aperture ovate, crenulated.” Measurements (mm.)— length width height 40.0 30.0 12.0 (Drouet) Synonymy— 1858 Patella moreleti Drouet, Moll. Mar. Agores, p. 42, pli 2, figs. 10, 11. 1891 Patella moreleti Drouet, Pilsbry, Man. Conch., vol. 13, p. 85, pl. 56, figs. 27, 25. [01-583] 100 Subgenus Patella Plate 71. Patella piperata Gould, 1846. Madeira, 27 mm., AWBP coll. 1058. Patella lowei Orbigny, 1839 (Pl. 74, figs. 1, 2) Range—Canary Islands. Remarks—This species appears to be closely allied to the Mediterranean caerulea, from which it differs mainly in having stronger, and more regu- lar sculpture, resulting in a more even denticula- tion of the margin, as opposed to the 6 or 7 distinct marginal angles of caerulea. Also, the coloration in lowei is darker, the exterior being rusty-brown, and the interior dark bluish to reddish brown at the edges, reflecting iridescent blue, and always with a clearcut white spatula. Description—Shell of moderate size, up to 56.5 mm. (24 inches) in length, ovate, depressed, with the apex towards the anterior third, solid but not thick, densely sculptured with broadly rounded primary radials and narrow intermediates. The margin is strongly and regularly corrugated, the projections compound and foliated. Colour as described above. Measurements (mm.)—(both A. W. B. Powell col- lection). very length width height 56.5 45.0 13.0 ‘Teneriffe 54.5 44.5 11.0 Teneriffe Synonymy— 1839 Patella lowei Orbigny, in Webb and Berthelot, Hist. Nat. Canaries, Moll., vol. 2, p. 97, pl. 7, figs. 9, 10. 1839 Patella azorica Nuttall, in Jay, Cat. Shells, ed. 3, p. 38. 1S91 Patella caerulea var. lowei Orbigny, Pilsbry, Man. Conch., vol. 13, p. 84, pl. 29, figs. 44-46; pl. 53, figs. 7-11. Records—CANARY ISLANDS (Orbigny); Teneriffe (AWBP coll. 5268). A. W. B. Powell Patellidae Plate 72. Figs. 1, 2. Patella caerulea Linnaeus, 1758, Isle of Capri, Italy, 51 mm., AWBP coll. 211. Figs. 3, 4. Patella safiana Lamarck, 1819, Oran, Algeria, 65 mm., AWBP coll. 1959. Patella gomesii Drouet, 1858 (PI. 74, figs. 5, 6) Range—Azores, Bay of San Lourenzo, Santa Maria and Pico. Remarks—The writer has not seen examples of this shell, which may prove to be a form of lowei. Its distinctive character is in having about 14 very prominent rounded radial folds, the whole surface, folds included, being densely radially lirate. Description—(Pilsbry’s translation of original): “Shell large, subdepressed, rugose, ribbed-plicate, rather solid; outside grayish-brown or rufescent; inside shining, brown, pearly; apex situated at the front third of the length, obtuse; aperture oval, entire.” Measurements (mm.)— length width height 50-60 50-53 12-15 Synonymy— 1858 Patella gomesii Drouet, Moll. Mar. Iles Agores, p. 39, pl. 1, figs. 6, 7. 1891 Patella gomesii Drouet, Pilsbry, Man. Conch., vol. 13, p. 86, pl. 54, figs. 17, 18. [01-584] November 27, 1973 Plate 73. Antarctic Australian Cellana and INDO PACIFIC MOLLUSCA, vol. 1, no. 15 Patella 101 (for plate 70, see p. 105) Subantarctic Nacella Figs. 1-3. Cellana framoserica (Holten, 1802). Fig. 1. South Australia. Fig. 2. Caloundra, Queensland. Fig. 3. Torquay, Victoria. Figs. 4-6. Cellana solida (Blainville, 1825). Figs. 4, 5. Stan- ey, Tasmania. Fig. 6. South Australia (rubraurantiaca orm). Figs. 7, 8. Cellana ardosiaea (Hombron & Jacquinot, 1841). Island of Juan Fernandez. Fig. 9. Nacella mytilina (Helbling, 1779). Falkland Islands. Fig. 10. Nacella kerguelenensis (E. A. Smith, 1877). Heard Island. 7 - Fig. 11. Nacella (Patinigera) deaurata (Gmelin, 1791). Falk- land Islands. Fig. 12. Nacella (Patinigera) terroris (Filhol, 1880). Campbell Island. a Fig. 13. Nacella (Patinigera) clypeater (Lesson, 1831). Chile. Figs. 14, 15. Nacella (Patinigera) magellanica (Gmelin, 1791). Possession Bay, Patagonia. [01-585] 102. Subgenus Patella A. W. B. Powell Patellidae [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence. | [01-586] November 27, 1973 Subgenus Uncertain The following six species of Patella are insuffi- ciently understood, particularly with regards to their soft anatomy, to be assigned as yet to their proper subgenera. Patella candei Orbigny, 1839 (PI. 75, figs. 7, 8) Range—Canary Islands. Remarks—The writer has not seen this species, but from published information it appears to be closely allied to citrullus from Funchal. Madeira. The surface has subobsolete radials crossed by prominent wavy concentric lirations that hap- hazardly anastomose to form an irregular netted appearance. Description—(Pilsbry’s 1891 translation of the original description): “Shell elevated, conical, thick, smooth or irregularly roughened; ovate, margin en- tire. Inside buff, bluish in the middle, outside pale yellow.” Measurements (mm.)— length width height 67.0 55.0 27.0 Synonymy— 1839 Patella candei Orbigny, in Webb and Berthelot, Hist. Nat. Canaries, vol. 2, Moll., p. 98, pl. 7, figs. 11, 12. 1854 Patella candei Orbigny, Reeve, Conch. Iconica, vol. 8, pl. 15, figs. 34 a, b. 1891 Patella candei Orbigny, Pilsbry, Man. Conch., vol. 13, p. 86, pl. 55, figs. 22-24. Patella citrullus Gould, 1846 (Pl. 75, figs. 9, 10) Range—Funchal, Madeira. Remarks—The writer has not seen examples of this species which appears to be related to candei. Pilsbry (1891, l.c.) remarked that the external sur- face resembles the skin of a cucumber. Description— (original) “Shell sub-diaphanous, thin sub-conical, moderately elevated, summit prominent; apex anterior, acute, feebly incurved, usually somewhat eroded; a great number of faintly INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella = 103 elevated lines, studded with fine tubercles or asper- ities, radiate from it, and become obsolete about half way towards the margin. Striae of increment coarse and irregular, overlaying each other, so as to give the shell a rude, concentrically squamose aspect externally; disc nearly oval, a little narrowed anteriorly; margin very thin and sharp, finely and irregularly undulated. External colour a dusky olive-green, with a shade of brown showing through it, ornamented with concentric, undulating lines of obscure white. Interior greenish-white, with bright iridescent reflections; a slight spatulaform deposit at the fundus, bluish at the edges and forepart, passing into greenish towards the middle and pos- terior portions.” Measurements (mm.)— length width height 45.0 32.0 7.0 Synonymy— 1846 Patella citrullus Gould, Proc. Boston Soc. Nat. Hist., vol. 2, p. 149. 1891 Patella citrullus Gould, Pilsbry, Man. Conch., vol. 13, p. 86, pl. 28, figs. 39-41. 1964 Patella citrullus Gould, Johnson, U. 239, p. 56. S. Nat. Mus. Bull. Patella concolor Krauss, 1848 (Pls. 64, 76, 78) Range—Natal coast to as far south and west as Bushman’s River, near Port Elizabeth. Remarks—The former name of this well-known South African limpet, Patella variabilis Krauss, 1848, is invalidated by two prior homonyms, those of Roding, 1798, and Risso, 1826. The earliest valid name to replace variabilis is concolor which is the uniformly dark-ashen colour form of this species. The species is exceedingly variable in colour pat- tern, but the shape, which is ovate, distinctly nar- rowed in front, remains constant. Also it is of light build and is often semi-transparent. Description—Shell rather small, usually between 30 and 35 mm. in length, but occasionally attaining 50 mm. (2 inches) in length, of light build, some- times semi-transparent, rather depressed, ovate, but distinctly narrowed at the anterior end. Sculptured with about 80 fine but somewhat unequal radial ribs, crossed by dense inconspicuous concentric lirae. Apex subcentral to about the anterior third, the area in its vicinity usually smooth. Colour ex- extremely variable, ranging from plain yellow, pale yellowish brown, and rusty-brown (concolor) to al- most black, and variously maculated; sometimes the [01-597] 104. Subgenus uncertain A. W. B. Powell Patellidae yellow form has one, or several, dark-brown radial streaks, and the black form (polygramma) has the primary radials picked out in white; the common form is pale yellowish brown, radially lined and speckled in dark-brown; spatula ill-defined, light brownish or clouded with white callus. Radula—Formula 3 + 1 + (2+0+2) + 1 + 3. The radula differs from those of all other South African patellids in the absence of the middle member of the central teeth; the four remaining centrals, how- ever, are grouped in pairs with a space between them, whereas in the radula of Patella vulgata and other European patellids, the four centrals are closely grouped, without space for a middle mem- ber. Plate 74. Figs. 1, 2. Patella lowei dOrbigny, 1839, Teneriffe, Canary Islands, 56.5 mm., AWBP coll. 5268. Figs. 3, 4. Pa- tella ct. caerulea Linnaeus, 1758 (crenata form), Madeira, 70 mm., AWBP coll. 675. Figs. 5, 6. Patella gomesii Drouet, 1858, Azores, 50-60 mm. From Pilsbry, 1591, pl. 54, figs. 17, 18. Measurements (mm.)— length width height 50.0 45.0 11.25 South Africa 48.5 41.25 14.00 Natal coast 35.0 30.5 9.00 Port Alfred 28.5 24.0 7.00 Coffee Bay Synonymy— 1848 Patella variabilis Krauss, Sudafr. Moll., Stuttgart, p. 55, pl. 3, fig. 12 (non P. variabilis Risso, 1826). 1848 Patella variabilis var. fasciata Krauss, Sudafr. Moll., Stutt- gart, p. 55, pl. 3, fig. 12 a (non P. fasciata Gmelin, 1791). 1848 Patella variabilis var. radiata Krauss, Sudafr. Moll., Stutt- gart, p. 59, pl. 3, fig. 12 b. (non P. radiata Born, 1778). 1548 Patella variabilis var. concolor Krauss, Sudafr. Moll., Stuttgart, p. 55, pl. 3, fig. 12 c. 1891 Helcioniscus variabilis Krauss, Pilsbry, Man. Conch., vol. 13, p. 147, pl. 16, figs. 18-20. Plate 75. Figs. 1, 2. Patella baudonii Drouet, 1858, Azores, 60 mm. Figs. 3, 4. Patella moreleti Drouet, 1858, Fayal, Azores, 40 mm. Figs. 5, 6. Patella rangiana Rochebrune, 1882, Cape Verde Islands, 44 mm. Figs. 7, 8. Patella candei d@Orbigny, 1839, Canary Islands, 67 mm. Figs. 9, 10. Patella citrullus Gould, 1846, Funchal, Madeira, 45 mm. (All figures from Pilsbry, 1891, Manual of Conchology, vol. 13, plates 45, 54, 55, 56 and 58). [01-598] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 1, no. 15 Patella 105 Plate 70. New Zealand Cellana Figs. 1-4. Cellana radians (Gmelin, 1791). Fig. 1. Mount Maunganui, Bay of Plenty (earlii pattern). Figs. 2, 3. Mo- tuihi Island, Auckland. Fig. 4. Herekopare Island, Stewart Island (perana form). Figs. 5, 6. Cellana flava (Hutton, 1873). Fig. 5. East Cape. Fig. 6. Cape Campbell, Marlborough. Figs. 7, 8. Cellana denticulata (Martyn, 1784). Mount Maun- ganui, Bay of Plenty. Figs. 9-11. Cellana stellifera (Gmelin, 1791). Fig. 9. Whan- (for plate 73, see p. 101) garei Heads. Fig. 10. Ti Point, Hauraki Gulf. Fig. 11. Long Beach, Bay of Islands (bleached coloration of beach shells). Figs. 12, 13. Cellana ornata (Dillwyn, 1817). Fig. 12. Mo- tutara, West Coast, Auckland. Fig. 13. Mount Maunganui. Figs. 14-16. Cellana strigilis (Hombron & Jacquinot, 1841). Campbell Island. Figs. 17-19. Cellana strigilis subspecies redimiculum (Reeve, 1854). Kartigi Beach, North Otago, South Island, New Zealand. [01-599] 106 Subgenus uncertain A. W. B. Powell Patellidae 1921 Patella variabilis constellata G. B. Sowerby, Proc. Malac. Soc., Lond., vol. 14, p. 127. 1931 Patella variabilis Krauss, Tomlin, Ann. Natal Mus., vol. 6 (3), p. 417. 1931 Patella variabilis fasciolata Tomlin, Ann. Natal Mus., vol. 6 (3), p. 418; nom. nov. pro P. variabilis fasciata Krauss, 1848 (non Gmelin, 1791). 1931 Patella variabilis polygramma Tomlin, Ann. Natal Mus., vol. 6 (3), p. 418; nom. nov. pro P. variabilis radiata Krauss, 1848 (non Born. 1778). 1932 Patella variabilis Krauss, Turton, Mar. Shells Port Alfred, p. 167, sp. 1187. 1932 Patella variabilis fasciata Krauss, Turton, Mar. Shells. Port Alfred, p. 167, sp. 1188. 1932 Patella variabilis radiata Krauss, Turton, Mar. Shells Port Alfred, p. 167, sp. 1189. 1932 Patella variabilis concolor Krauss, Turton, Mar. Shells Port Alfred, p. 168, sp. 1190. 1932 Patella variabilis constellata Sby., Turton, Mar. Shells Port Alfred, p. 168, sp. 1191. 1932 Patella variabilis helvola Turton, Mar. Shells Port Alfred, p. 168, sp. 1192. 1932 Patella rietensis Turton, Mar. Shells Port Alfred, p. 167, pl. 38, fig. 1183. 1932 Patella rota (non Gmelin, 1791) Turton, Mar. Shells Port Alfred, p. 168, sp. 1193. 1932 Patella helena Turton, Mar. Shells Port Alfred, p. 168, pl. 39, fig. 1194. 1932 Patella conspicua (non Philippi, 1849) Turton, Mar. Shells Port Alfred, p. 168, sp. 1196 (in part). 1932 Patella farquhari Turton, Mar. Shells Port Alfred, p. 170, pl. 40, fig. 1207. 1949 Patella variabilis Krauss, Koch, Ann. Natal Mus., vol. 11 (3), p. 510, pl. 23, figs. 1-11; text figs. 21, 22 (radula). Records—SOUTH AFRICA: Natal coast to as far south and west as Port Elizabeth (Koch, 1949); Natal (ex Koch; AWBP coll.); Umtwalumi, 22 miles N. of Port Shepstone (V. Orr, 1955; ANSP); Port St. John’s Pondoland (V. Orr; ANSP); Coffee Bay, Transkei (V. Orr, 1955; ANSP); Port Alfred (USNM); (AWBP coll.); near Durban (Mrs. N. Prior). Patella depsta Reeve, 1855 (PI. 77; pl. 78, fig. 1) Range—Islands of St. Paul and Amsterdam, South Indian Ocean. Remarks—Reeve cited “Macao and the Island of St. Paul” as localities for this species, but the first mentioned location is obviously a mistake. St. Paul is here nominated as the type locality. The species also occurs at the adjacent island of Amsterdam. Gaillard (1954) figured the radula of depsta, and assigned the species to Cellana, but the radula sug- gests much closer alliance with the Patellinae, and except for the laterals, is not unlike that of Patella (Patellona). The laterals in. the Patellinae are usu- ally fused at the base, and have a pluricuspid head, but Gaillard’s drawing shows a pair of laterals on either side, each separated at the base. Since the writer has no preserved material of this species the Plate 76. Figs. 1-7. Patella concolor Krauss, 1548. Figs. 1, 2. Port Alfred, South Africa, 35-36 mm., AWBP. coll. 30872; 227788. Figs. 3, 4. Port St. Johns, Pondoland, South Africa, 26-34 mm., AWBP coll. 30822. Fig. 5. Coffee Bay, Transkei, South Africa, 30 mm., AWBP coll. 48225. Fig. 6. Umtwalumi, South Africa, 34 mm., AWBP coll. 211735. Fig. 7. South Africa, 50 mm., AWBP coll. 46130. [01-600] November 27, 1973 Plate 77. Patella depsta Reeve, 1854. Island of St. Paul, South Indian Ocean, 32-35 mm., AWBP coll. 46133. apparently unusual form of the laterals cannot be confirmed at present. Description—Shell of moderate size, up to 35.5 mm. (1% inches) in length, lightly built, ovate, gradually narrowed in front, and moderately ele- vated, with the apex anterior to the middle, com- pressed and hooked. Sculptured finely and deli- cately radially lirate, arranged more or less in fours, the inner two weaker than the outer two, and about 100 lirae in all. Colour of exterior pale pinkish chestnut; interior orange-brown with a slight bronzy sheen; spatula pinkish white. Plate 78. Fig. 1. Patella depsta Reeve, Island of St. Paul. Radula, from Gaillard, 1954, p. 521, fig. 1. Fig. 2. Patella concolor Krauss, Natal. Radula, from Koch, 1949, p. 511, fig. 22 (as variabilis Krauss). INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 107 Measurements (mm.)—(both A. W. B. Powell collection). length width height 35.5 28.0 14.0 St. Paul 32.25 24.0 12.0 St. Paul Radula—Formula 3 + 2? + (2+1+2) + 2? + 3. Synonymy— 1855 Patella depsta Reeve, Conch. Iconica, pl. 31, figs. 85 a, b. (Jan.). 1891 Helcioniscus depsta Reeve, Pilsbry, Man. Conch., vol. 13, p. 151, pl. 20, figs. 45, 46. 1954 Cellana devsta Reeve, Gaillard, Bull. Mus. Nat. d’Hist. Nat., vol. 26, pp. 520, 521, text. fig. 1 (radula). Patella rangiana Rochebrune, 1882 (Pl. 75, figs. 5, 6) Range—Porto Praya, Cape Verde Islands. Remarks—This species, which the writer has not seen, appears to be a distinctive one, with its very scaly prominent radial ribs. Pilsbry’s translation of the original description follows, accompanied by copies of Rochebrune’s figures. Description—‘“Shell ovate, depressed-convex, rufous; vertex submucronate, usually eroded, situated at 2/3 of the length; having larger and smaller radiating broad, very scaly ribs, scales subimbricating, obtuse, lenticular; margin un- dulating; interior bluish, silvery-pearly, rayed with bands and spots of purplish, the center spatulate.” Measurements (mm.)— length width height 44.0 36.0 19.0 (Rochebrune) Synonymy— 1882 Patella rangiana Rochebrune, Bull. Soc. Philomathique, Paris, ser. 7, vol. 6, p. 29. 1891 Patella rangiana Rochebrune, Pilsbry, Man. of Conch., vol. 13, p. 89, pl. 58, figs. 42, 43. ? Patella kaffraria Rennie, 1930 (PI. 79) Range— Upper Cretaceous of Pondoland, South Africa. Remarks—The author of this species remarked that “It need hardly be stated that the genus Patella is here used in the widest possible sense. The species is apparently distinct from any pre- viously described from the Cretaceous.” It certainly appears to belong to the Patellacea, but a precise generic or even familial allocation would be purely conjectural. Rennie’s original description follows. [01-601] 108 Subgenus uncertain Plate 79. PPatella kaffraria Rennie, 1930. Upper Cretaceous of Pondoland, South Africa, 32.56 mm. Holotype, from Rennie, 1930, pl. 24, figs. 1, 2. Description—“Shell moderately convex, with the apex obtusely pointed, not recurved, and placed well in front of the middle; the sides are straight, or only slightly convex. Aperture oval, A. W. B. Powell Patellidae considerably longer than wide, with wavy margin. Surface with stout, rather irregular, radial ribs, and narrow furrows; on the posterior side the ribs are of two sizes, the larger and smaller alternat- ing; on the anterior side there are more numer- ous, finer ribs; the ribs are crossed by irregular growth markings.” Measurements (mm.) (Not stated, but evidently the figures are natural size)— length width height 32.5 25.0 14.0 holotype Synonymy— 1930 Patella kaffraria Rennie, Annals of South African Mu- seum, vol. 28, p. 206, pl. 24, figs. 1-4. Types— The holotype (No. 8477) and paratype (No. 8572) are in the South African Museum. (01-602] November 27, 1973 Subgenus Patellona Thiele in Troschel, 1891 Type: Patella granatina Linnaeus, 1758 This group of patellids was named because of a marked difference in the radula from that of typ- ical Patella. The four central teeth of true Patella occur in a horizontal alignment whereas in Patel- lona there is a median central, narrower and of smaller size than the outer pairs of centrals, which instead of being in line, descend steeply to the laterals, their tops forming a chevron. The cusps of the centrals and laterals vary between oblique heart-shape and parrot-beaklike. This chevron-like radula is found in species from Cape Verde Islands, Senegal, Guinea, An- gola and St. Helena, as well as South Africa, where the type species granatina and the related oculus occur, these two being more or less re- stricted to the cooler waters of the west coast. Shells of this subgenus are slightly iridescent within and the shell substance is sufficiently transparent for the external colour patterns to show through faintly. The northward flowing cool Benguela Current could account for the presence of the subgenus in Angola and St. Helena, but locations north of there, in the tropical waters of West Africa and the Cape Verde Islands, are, under present con- ditions, out of range of the influence of that cur- rent. Nevertheless the style of radula in the trop- ical West African and Cape Verde Islands limpets is so similar to that of the cool water species of the South African west coast that some distribu- tional continuity, under more uniform hydro- logical conditions, must have existed in the past. Related to Patellona is the subgenus Cymbula (see ahead) in which the central teeth have the same chevronlike alignment, but their cusps are distinctive in having broad blunt tops with raised marginal rims. Synonymy— 1891 Patellona Thiele in Troschel, Das Gebiss der Schnecken, vol. 2, p. 317, for granatina Linnaeus, 1758, adansonii Dunker, 1853 and plumbea Lamarck, 1819. Type, by subsequent designation, Tomlin, 1931: Patella grana- tina Linnaeus, 1758S. INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella = 109 Patella granatina Linnaeus, 1758 (Pl. 64, figs. 1-3; pl. 80; pl. 82, fig. 2) Range—South Africa, Port Nolloth on the west coast, south to False Bay and extending east- ward to Danger Point. Remarks—This large but relatively thin South African limpet is easily recognised by its broadly ovate, almost pentagonal outline, strong, nar- rowly crested, radial ribs, and distinctive colora- tion of the interior which is bluish white with a clearly outlined dark-brown spatula. Description—Shell large, up to 85 mm. (3% inches) in length, strong but of relatively light build, broadly ovate and _ tall-conical, with the apex almost central. Sculpture of radiate folds that strongly corrugate the margin; five of the radials on the posterior half of the shell are stronger than the rest; radials and interspace alike bear closely-spaced cords that are rendered scabrous by dense concentric growth-lamellae. Colour of exterior greyish to dull-white with an underlying pattern of dark-brown, zigzag, con- centric markings, often forming a netted design; interior bluish white, the spatula dark-chocolate, with clearly defined edges, and a marginal pat- tern of numerous short, dark-brown dashes, with regular gaps corresponding to the external pri- mary radials. Radula—Formula 3+ 1 + (2+1+2) + 1+ 3. The median central is small and slender, flanked by a pair of stout fully-developed centrals on either side, followed by a pluricuspid lateral, and the usual three, more or less functionless, marginals. The centrals, collectively, form a chevron, as in other members of this subgenus. The cusps of the paired centrals and the pluricuspid laterals are leaf-shaped, obliquely flexed, and with a median groove or depression. Plate 80. Patella (Patellona) granatina Linnaeus, 1758. Table Bay, South Africa, 71 mm., AWBP coll. 183. [01-613] 110 Subgenus Patellona A. W. B. Powell Patellidae Measurements (mm.)—(all A. W. B. Powell collection) length width height 85.0 78.5 30.0 South Africa 74.0 63.5 35.0 South Africa 60.0 49.5 20.0 False Bay Synonymy— 1758 Patella granatina Linnaeus, Syst. Nat. ed. 10, p. 782. 1819 Patella apicina Lamarck, Anim. sans vert., vol. 6 (1), p. 324. 1848 Patella granatina Lam., Krauss, Sudafr. Moll., Stuttgart, p. 43. 1854 Patella granatina Lam., Reeve, Conch. Iconica, pl. 3, figs. 4a, b. 1891 Patella (Scutellastra) granatina Lam., Pilsbry, Man. Conch., vol. 13, p. 106, pl. 62, figs. 76, 77. 1891 Patellona granatina Lam., Thiele, Das Gebiss der Schnecken, vol. 2, p. 317. 1931 Patellona granatina Lam., Tomlin, Ann. Natal Mus., vol. 6 (3), p. 417 (designated type of Patellona) 1949 Patella granatina Lam., Koch, Ann. Natal Mus., vol. 11 (3), p. 501, pl. 20, figs. 1-5; text figs. 9, 10 (radula) Records—SOUTH AFRICA: Table Bay (AWBP coll.); False Bay (AWBP coll.); Platboom, Cape Peninsula (V. Orr, 1955; ANSP); Sea Point (Mrs. N. Prior) Simonstown (AWBP coll.). Patella oculus Born, 1778 (PI. 64, figs. 7-9; pl. 81; pl. 82, fig. 1) Range—South Africa, west coast from near Cape Town eastward to Umhlali. Remarks—This large, depressed, broadly-ovate, star-shaped limpet has something of the appear- ance of Patella (Scutellastra) longicosta Lam- arck, but differs from it, not only in dentition, but also in coloration, for the interior of oculus is dark purplish brown, except for a yellowish brown spatula and a surrounding area of light bluish grey. The species is essentially a cold-water one, and is more abundant along the west coast of South Africa than to the eastward. It occurs in the Bala- noid zone, which is lower mid-tidal, but sometimes extends to and below low spring-tide level. Description—Shell large, up to 110 mm. (4% inches) in length, solid, depressed, broadly ovate, star-shaped, with the principal ribs strongly cor- rugating the margin. Sculpture consisting of about 11 primary, broad, carinated radials, plus secondary radials and interstitial threads. Colour of exterior dull-brown to blackish, but usually eroded to dull-light greyish brown; interior with a very broad dark purplish brown border, and a light bluish grey area surrounding the spatula, which is fawn to deep yellowish brown. Radula—Formula 3 + | + (2+1+2) + 1 + 3, very similar to the radula of granatina, the central teeth having the same chevronlike alignment. Measurements (mm.)— length width height 110.0 106.0 42.0 Cape of Good Hope 86.0 76.0 15.0 Port Elizabeth 73.0 63.0 15.0 Port Alfred 53.0 51.0 8.0 Still Bay Plate 81. Patella (Patellona) oculus Born, 1778. Cape of Good Hope, South Africa, 77 mm., AWBP coll. 200. [01-614] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 111 Plate 82. Fig. 1. Patella (Patellona) oculus Born, South Africa. Radula, from Koch, 1949, p. 508, fig. 18. Fig. 2 Patella (Pa- tellona) granatina Linnaeus, South Africa. Radula, from Koch, 1949, p. 502, fig. 10. Fig. 3. Patella (Patellona) canescens Gmelin, St. Helena. Radula, from Thiele, 1891, pl. 28, fig. 7 (as plumbea Lamarck). Fig. 4. Patella (Patellona) adansonii Dunker, “Chinchao” in error, probably West Africa. Radula, from Thiele, 1891, pl. 28, fig. 8. Synonymy— 1778 Patella oculus Born, Index Mus. Caes. Vind., p. 434. 1786 Patella oculus hirci Lightfoot, Cat. Portland Mus., p. 105. 1791 Patella badia Gmelin, Syst. Nat., ed. 13, p. 3700. 1791 Patella monopis Gmelin, Syst. Nat., ed. 13, p. 3707. 1791 Patella fuscescens Gmelin, Syst. Nat., ed. 13, p. 3701. 1798 Patella astrolepas Réding, Mus. Bolten., vol. 2, p. 12. 1819 Patella scutellaris Lamarck, Anim. sans vert., vol. 6 (1), p. 328. 1848 Patella schroeteri Krauss, Siidafr. Moll., Stuttgart, p. 43. 1854 Patella oculus Born, Reeve, Conch. Iconica, pl. 2, figs. 2a, b. 1891 Patella (Scutellastra) oculus Born, Pilsbry, Man. Conch., vol. 13, p. 106, pl. 27, figs. 30-32. 1932 Patella oculus Born, Turton, Mar. Shells Port Alfred, p. 162. 1932 Patella oculus badia Gmelin, Turton, Mar. Shells Port Alfred, p. 163. 1932 Patella oculus fuscescens Gmelin, Turton, Mar. Shells Port Alfred, p. 163. 1932 Patella oculus schroeteri Krauss, Turton, Mar. Shells Port Alfred, p. 163. 1932 Patella oculus planulata Turton, Mar. Shells Port Al- fred, p. 163. 1942 Patella oculus Born, Tomlin & Stephenson, Proc. Malac. Soc., Lond., vol. 25, pp. 5, 6. 1949 Patella oculus Born, Koch, Ann. Natal Mus., vol. 11 (3), p. 507, pl. 22, figs. 1-4; text figs. 17, 18 (radula). 1967 Patella scutellaris Lamarck, Christiaens, Bull. Mus. Nat. d’Hist. Nat. ser. 2, vol. 39 (5), p. 973. Records—SOUTH AFRICA: west coast near Cape Town, eastward to Umhlali (Koch, 1949, p. 507); Cape of Good Hope (AWBP coll.);False Bay (AWBP coll.); Buffel’s Bay, Cape Pen- insula (Mrs. N. Prior); Still Bay (Auck. Mus.); Port Elizabeth (Auck. Mus.); Jeffrey's Bay (Auck. Mus.); Port Alfred (AWBP coll.). Patella adansonii Dunker, 1853 (Pl. 82, fig. 4; PI. 83, figs. 5, 6) Range—West Africa. Remarks—This species is characterised by its dense fine radial ribbing, brown-lined and mar- bled external pattern, and finely denticulated margin. Description—Shell of moderate size, 36-50 mm. (1%-2 inches) in length, ovate, the anterior end slightly narrowed, moderately elevated, with the apex at about the anterior third. Sculpture con- sisting of about 80 to 100 narrowly-rounded ra- dial ribs that more or less alternate in strength. Colour, externally whitish, marbled, and radially and narrowly streaked with olive or dark greenish brown, internally pale bluish grey, with the ex- ternal pattern showing through, more strongly at the margin; spatula buff to pale orange-brown. Radula—Formula 3 + 1 + (2+1+2) + 1+ 3. The five central teeth are not in a straight horizontal row as in typical Patella, for the outer pair of centrals are lower than the inner pair, and the median one is small, very slender and almost vestigial. Measurements (mm.)— length width height 50.0 41.0 17.0 42.0) 33.0 14.0 Pilsbry, 1891, p. 92 Angola [01-615] 112. Subgenus Patellona A. W. B. Powell Synonymy— 1853 Patella adansonii Dunker, Ind. Moll. Guin. Infer., p. 42, pl. 6, figs. 10-15. 1891 Patella adansonii Dunker, Pilsbry, Man. Conch., vol. 13, p. 92, pl. 12, figs. 30-33. Records—WEST AFRICA: Loanda (type); Ambrizette, An- gola (AWBP coll.; ANSP). Patella canescens Gmelin, 1791 (Pl. 82, fig. 3; Pl. 83, figs. 3, 4) Range—St. Helena. Remarks—The sculpture is much finer than that in either lugubris or plumbea and in conse- quence the shell margin is delicately crenulated rather than corrugated. Plate 83. Figs. 1, 2. Patella (Patellona) lugubris Gmelin, 1791. Porto Grande, St. Vincent, Cape Verde Islands, 42-54 mm., AWBP. coll. 50089. Figs. 3,4. Patella (Patellona) canescens Gmelin, 1791. St. Helena, 41-45 mm., AWBP coll. 125412. Figs. 5, 6. Patella (Patellona). adansonit Dunker, 1553, Anvola, West Africa, 36-42 mim., AWBP coll. 146139. Patellidae Description—Shell of moderate size, up to 47 mm. (1% inches) in length, ovate, slightly nar- rowed in front, moderately elevated, with the apex subcentral. Sculpture crisp, consisting of very numerous radial cords, the primaries grouped in pairs or in threes, with an occasional intermediate between each group of primaries. Colour of exterior black, usually eroded to a greyish brown. Interior bluish silvery, the spat- ula flesh to orange-brown, and the edge of the shell narrowly margined in black. Measurements (mm.)— length width height 47.0 40.0 20.0 St. Helena; Christiaens, 1968 45.0 37.5 20.0 St. Helena Radula—F ormula 3 + 1 + (2+1+2) + 1 +3. Radula very similar to that of plumbea and adansonii, with the median central very small and the pairs of centrals arranged chevron-like, descending sharply from the central line. The lateral is dis- tinctive in having four well developed cusps (Christiaens, 1968, text fig. 1). Synonymy— 1791 Patella canescens Gmelin, Syst. Nat., ed. 13, p. 3724. Locality? 1855 Patella canescens Gmelin, Reeve, Conch. Iconica, pl. 34, figs. 103 a, b. Locality? 1968 Patella canescens Gmelin, Christiaens, Rev. Zool. Bot. Afr., vol. 77, pts. 3-4, pp. 314-320. St. Helena. Patella lugubris Gmelin, 1791 (PI. 83, figs. 1, 2) Range—West Africa, Loanda, Benguela, Guinea and Cape Verde Islands. Remarks—The species is much more coarsely ribbed than either the St. Helena canescens or the West African plumbea, and from the latter species it differs in being more broadly ovate. Description—Shell moderately large, up to 60 mm. (2% inches) in length, broadly ovate, mod- erately elevated, with the apex almost at the anterior third. Sculpture consisting of numerous strong, keeled radials that prominently corrugate the margin. Colour, externally dull-black, inter- nally silvery bluish grey, the spatula often clouded with a white callus. Measurements (mm.)— length width height 60.0 50.0 20.0 Pilsbry, 1891, p. 91 53.0 46.0 19.0 Cape Verde Islands [01-616] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, No. 15 Patella 113 Synonymy— 1791 Patella lugubris Gmelin, Syst. Nat. ed. 13, p. 3705; based upon Martini-Chemnitz, Conch. Cab., vol. 1, pl. 8, fig. 60. 1854 Patella lugubris Gmelin, Reeve, Conch. Iconica, pl. 14, figs. 32 a-c. “Island of St. Vincent, West Indies,” sic. - St. Vincent, Cape Verde Islands. 1891 Patella lugubris Gmelin, Pilsbry, Man. Conch., vol. 13, p. 90, pl. 12, figs. 39, 40, 41-44; pl. 57, figs. 32-35. Patella plumbea Lamarck, 1819 (PI. 84) Range—Senegal, West Africa. Remarks—This species is closely allied to lugu- bris, and when adequate material is studied, may prove to be identical. Pilsbry (1. c.) remarked that “the ribbing is finer than in P. lugubris, the shell is more elliptical, more depressed, and the central spatula of the interior is longer and nar- rower.” Description—Shell moderately large, up to 53 mm. (2-1/16 inches) in length, narrowly ovate, low-conical. Colour, externally dull-black, inter- nally bluish, the spatula whitish, often clouded with brown. Measurements (mm.)— length width height 52.0 38.0 10.5. type; Mermod, 1950, p. 692 Synonymy— 1819 Patella plumbea Lamarck, Anim. s. Vert., vol. 6, p. 328. 1834 Patella caerulea Quoy and Gaimard, Voy. Astrolabe, Moll., vol. 3, p. 342, pl. 70, figs. 4-6. 1854 Patella plumbea Lam., Reeve, Conch. Iconica, pl. 3, figs. 3a, b. Plate 84. Patella (Patellona) plumbea Lamarck, 1819. Senegal, West Africa. Holotype, 52 mm., from Mermod, Rev. Suisse Zool., vol. 57, no. 34, p. 693, fig. 4. 1891 Patella plumbea Lam., Pilsbry, Man. Conch., vol. 13, p. 91, pl. 24, figs. 11, 14, 15; pl. 57, figs. 38, 39. 1950 Patella plumbea Lam., Mermod. Rev. Suisse Zool., vol. 57, no. 34, pp. 692, 693, text fig. 4 (type). Patella safiana Lamarck, 1819 (PI. 72, figs. 3, 4) Range—Algeria, Morocco and West Africa. Remarks—This is a large, elongate-ovate spe- cies, only moderately elevated, and with a long narrow spatula. In coloration the exterior is whit- ish, with conspicuous brown rays in the rib inter- stices. The interior is silvery grey, with the brown external rays showing through towards the mar- gin; the spatula is creamy-white, often stained with orange-brown. Description—Shell large, up to 77 mm. (3 inches) in length, elongate-ovate, moderately ele- vated, with the apex at about the anterior third. Sculptured with broadly rounded primary radial ribs and weak interstitial cords. Colour: exter- nally with whitish primary ribs and the interstices intermittently rayed with brown; internally buff to silvery-grey, slightly iridescent; spatula cream, clouded with light orange-brown. Radula—F ormula 3 + 1 + 4 + 1 + 3. The radula resembles that of adansonii, canescens and plum- bea, in that the centrals are not in a horizontal line, the outer pair being lower than the inner pair. Also, a median central appears to be com- pletely absent, as in vulgata (see Fischer-Piette, 1935, p. 53, text fig. 22). Measurements (mm.)— length width height 77.0 57.0 21.0 largest of Lamarck’s type series 65.0 51.0 21.0 Oran, Algeria Synonymy— 1819 Patella safiana Lamarck, Anim. sans vert., vol. 6, p. 329. 1849 Patella conspicua Philippi, Abbild., vol. 3, p. 71. Guinea. 1852 Patella kraussii Dunker, Index Moll. Guin. inf., p. 42, pl. 6, figs. 4-6. 1854 Patella conspicua Philippi, Reeve, Conch. Iconica, pl. 7, fig. 12. Gaboon. 1891 Patella safiana Lam. Pilsbry, Man. Conch., vol. 13, p. 90, pl. 55, figs. 19-21. 1935 Patella safiana Lam. Fischer-Piette, Journ. Conchyl., vol. 79, p. 53. 1950 Patella safiana Lam., Mermod, Rev. Suisse Zool., vol. 57, no. 34, pp. 693, 694, text fig. 5. Types—The type series of safiana is in the Mu- séum D’Histoire Naturelle de Geneve. Records—ALGERIA: Oran (AWBP coll.). MOROCCO: ocean coast (Pilsbry, 1891). WEST AFRICA: Gaboon, Guinea (Phi- lippi, 1549, type of conspicua). [01-617] A. W. B. Powell Patellidae 114. Subgenus Patellona [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence. | [01-618] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 115 Subgenus Patellidea Thiele in Troschel, 1891 Type: Patella granularis Linnaeus, 1758 This subgenus, of which the type species is the only known member, appears to be most closely allied to the subgenus Scutellastra. The radula resembles that of Scutellastra in its main features, especially in having a narrow but well-developed median central, and the other 4 larger centrals in a horizontal line, but differs in the form of the cusps which are oblique and parrot-beaklike. The shell also differs from that of Scutellastra in texture in being more strongly coloured, and in having distinctive external sculpture, consisting of strong radials bearing prominent imbricated scales. Recent, South Africa. Synonymy— 1891 Patellidea Thiele in Troschel, Das Gebiss der Schnecken, volume 2, p. 315. Type, by monotypy: Patella granu- laris Linnaeus, 1758. Patella granularis Linnaeus, 1758 (pl. 64, figs. 4-6; pls. 85-87) Range—South Africa, the entire coastline from Port Nolloth in the west to Umpangazi in the east. Remarks—This common South African species is easily identified by its scaly external ribbing and bluish white interior, broadly margined in dark- brown, and with a reddish brown spatula. Reeve’s Patella vidua, erroneously recorded from the Philippines, is a synonym. A photograph of one of Reeve’s figured specimens (Fig. 22a) was kindly supplied by Dr. J. D. Taylor, and that specimen, in the collections of the British Museum (Natural History), is here nominated lectotype of vidua. Description—Shell of moderate size, up to 63 mm. (2% inches) in length, ovate, slightly narrowed in front, tall-conical, with the apex slightly anter- ior of the centre. Sculpture of strong, regular, rounded primary radial ribs, with slightly weaker intermediates; 1 to 3 intermediates between the primary radials; all ribs with closely-spaced, im- bricated, scales, resultant from numerous, lamel- lose concentric growth lines. Colour: externally dull light-brown to grey, with the scales paler; internally bluish white, with a dark-brown, wide, marginal border, and a reddish brown spatula. In fully grown examples the border is usually contin- uous, but in young shells it is interrupted by bluish white radial streaks, corresponding with the external radials. Radula—Formula 3 + 1 + (2+1+2) + 1+ 3. Central teeth 5, the middle member small and slender, almost vestigial, the outer pairs much larger, each with a long, pointed and incurved cusp; pluri- cuspid lateral with an enlarged top, bearing 4 cusps, of which the second from the proximal side is largest, and shaped like those of the central pairs; marginals 3, long, narrow and _ flexuous, each with a rudimentary cusp. Plate 85. Patella (Patellidea) granularis Linnaeus, 1758. Figs. 1, 2. Platboom, Cape Point, South Africa. Fig. 3. Cape of Good Hope, South Africa, 37-60 mm., AWBP coll. 42924 & 193. [01-629] 116 Subgenus Patellidae Measurements (mm.)— length width height 63.0 48.0 19.0 Sea Point; Mrs. N. Prior 59.0 49.0 26.0 C. of Good Hope 49.0 38.0 24.0 Platboom 44.5 36.0 17.0 Port Alfred Synonymy— 1758 Pateklla granularis Linnaeus, Syst. Nat., ed. 10, p. 782. 1834 Patella granularis L., Quoy and Gaimard, Voy. ‘Astro- labe’, Zool. vol. 3, p. 341, pl. 70, figs. 12-15. 1848 Patella. granularis’ L., Krauss, Sudafr. Moll., Stuttgart, p. 52. 1848 Patella echinulata Krauss, Sudafr. Moll., Stuttgart, p. 52, pl. 3, fig. 15. 1848 Patella natalensis Krauss, Sudafr. Moll., Stuttgart, p. 53, pl. 3, fig. 10. 1854 Patella vidua Reeve, Conch. Iconica, pl. 11, figs. 22a, b. 1854 Patella granularis L., Reeve, Conch. Iconica, pl. 14, figs. 31a, b. 1855 Patella morbida Reeve, Conch. Iconica, pl. 25, figs. 64a, b. 1891 Patella (Scutellastra) granularis L., Pilsbry, Man. Conch., vol. 13, p. 102, pl. 63, figs. 80-83. 1891 Patellidea granularis L., Thiele (new genus), in Troschel & Thiele, Das Gebiss der Schnecken, col. 2, p. 315. 1931 Patellidea granularis Linne, (designated type of genus) Tomlin, Ann. Natal. Mus., vol. 6 (3), p. 417. 1932 Patella granularis L., Turton, Mar. Shells Port Alfred, p. 166. Plate 86. Patella (Patellidea) granularis Linnaeus, 1758. Lec- totype, here nominated, of Patella vidua Reeve, 1854, er- roneously recorded from the Philippines, but considered to be synonymous with the South African granularis, The lecto- type is based upon Reeve’s fig. 22a. Photo by courtesy of Dr. J. D. Taylor, British Museum (Natural History). A. W. B. Powell Patellidae 1932 Patella morbida Reeve, Turton, Mar. Shells Port Alfred, p. 166. 1932 Patella natalensis Krauss, Turton, Mar. Shells Port Alfred, p. 166. 1932 Patella natalensis echinulata Krauss, Turton, Mar. Shells Port Alfred, p. 166. 1932 Patella miliaris Turton, Mar. Shells Port Alfred, p. 166. (non Philippi, 1848) 1932 Patella argenvillei assimilans Turton, Mar. Shells Port Alfred, p. 167. 1932 Patella alboradiata Turton, Mar. Shells Port Alfred, p. 167. 1933 Patella tomlini Turton, (nom. nov. pro P. alboradiata Turton, 1932, non Gmelin, 1791) Journ. Conch., vol. 19, p. 371. 1949 Patella granularis Linne Koch, Ann. Natal Mus., vol. II (3), p. 503, pl. 19, figs. 4-8; text figs. 11,12 (radula). Types—The types of granularis (Holotype; Sloane coll., no. 1013), morbida and vidua (lecto- type, here selected) are in the Britsh Museum (Natural History). Records—SOUTH AFRICA: Port Nolloth to Umpangazi, north of Durban (Koch, 1949, p. 503); Saldanha Bay (Dis- covery II, 1926); Table Bay (AWBP coll.); Cape of Good Hope (AWBP coll. 193); Sea Point (Mrs. N. Prior); False Bay (Auck. Mus.); Platboom, Cape Point (V. Orr, Jan. 1955); Jeffrey's Bay (AWBP coll.); Port Alfred (Auck. Mus.). K, | Plate 87. Fig. 1. Patella (Patellidea) granularis Linnaeus. South Africa. Radula. Fig. 2. Patella (Olana) cochlear Born. Radula, both from Koch, 1949, p. 504, fig. 12 & p. 499, fig. 6. [01-630] November 27, 1973 Subgenus Cymbula H. and A. Adams, 1854 Type (monotypy): Patella compressa Linnaeus, 1758 This subgenus contains two species, the shells of which are of very different outward appearance, but nevertheless have a striking sameness in the radula that is of a distinctive type. The multiple centrals and the pluricuspid lateral have large blunt-topped cusps, with strongly raised or flanged edges. The centrals collectively form a chevron instead of forming a horizontal line, as in typical Patella, and thus indicate alliance with the subgenus Patellona. The type species is the easily recognised com- pressa, with its elongated, laterally compressed shell, adapted to its specialised station on the stipes of the large kelps, Ecklonia and Laminaria. Only rarely is this species found attached to rock. On the other hand, the second species, miniata, is of normal ovate limpet shape, since it is ex- clusively a rock-dwelling. Both species are re- stricted to South African waters. Synonymy— 1854 Cymbula H. Adams and A. Adams, The Genera of Re- cent Mollusca, volume 1, p. 466. Type, by monotypy: Patella compressa Linnaeus, 1758. [Cymbula Gray, 1821, is an error for Cymbulia Peron and Lesnerr 1810]. Patella compressa Linnaeus, 1758 (pl. 63, fig. 9; Pls. 88, 89) Range—South Africa, from Port Nolloth in the west to Danger Point in the south. Records from further afield, including one from St. Helena, are due to drift, along with large algae, upon which the species lives. Description—Shell large, up to 117.5 mm. (4% inches) in length, thin, elongate-ovate, tall and narrow, with parallel sides, the apex a little for- ward of the middle, and curving anteriorly. Sculp- ture consisting of very numerous, rather unequal, linear-spaced riblets; margin very minutely crenu- lated, convex at the sides, and concave at the ends. Colour: externally dull brownish buff; inter- nally light pinkish fawn, the central area irregu- larly clouded with whitish callus. Radula—F ormula 3 + 1 + (2+1+2) + 1 + 3. Cen- tral teeth 5, forming a chevron, the median one INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 117 small and very narrow, bearing a simple small cusp, outer pairs of centrals massive, each with a broa,d flat-topped cusp, ridged on each side, the ridges more prominent on the outermost teeth; pluricuspid lateral with two cusps, each similar to those of the outer centrals; the three marginals are small, each with a single simple cusp, outer- most marginal largest of the three. Measurements (mm.)— length width height 117.5 51.0 50.0 Kommetje, Cape Peninsula; Mrs. N. Prior. 94.0 45.0 35.0 South Africa; Pilsbry, 1891, p. 93 83.0 44.0 36.5 South Africa 63.5 35.0 23.0 South Africa Synonymy— 1758 Patella compressa Linnaeus, Syst. Nat. ed. 10, p. 783. 1834 Patella compressa L., Quoy and Gaimard, Voy. ‘Astro- labe’, Zool., vol. 3, p. 338, pl. 70, figs. 1-3. 1848 Patella compressa L., Krauss, Siidafr. Moll., Stuttgart, p. 50. 1854 Patella compressa L., Reeve, Conch. Iconica, pl. 7, figs. 13a, b. 1854 Patella (Cymbula) compressa L., H. & A. Adams, Gen. Rec. Moll., vol. 1, p. 466. 1891 Patella compressa L., Pilsbry, Man. Conch., vol. 13, p. 93, pl. 61, figs. 68-70. 1949 Patella compressa Linne. Koch, Ann. Natal Mus., vol. 11, p. 499, pl. 17, figs. 4-6; text figs. 7,8 (radula). Records—SOUTH AFRICA: Port Nolloth to Danger Point (Koch, 1949, p. 499); Cape Peninsula (AWBP coll. 26039); Kommetje, Cape Peninsula (Mrs. N. Prior). Plate $8. Fig. 1. Patella (Cymbula) compressa Linnaeus. South Africa. Radula, from Koch, 1949, p. 500, fig. 8. Fig. 2. Patella (Cymbula) miniata Born. South Africa. Radula, from Koch, 1949, p. 507, fig. 16. [01-635] 118 = Subgenus Cymbula Patella miniata Born, 1778 (Pl. 63, figs. 10, 11; Pls. 88, 90, 91) Range—South Africa, from Port Nolloth in the west, eastward to Qolora, near East London, and Natal. Remarks—This moderately large, very attractive limpet is ovate, depressed and relatively thin, with an intricate pattern of radial streaks and speckles; it is reddish brown in living examples, but bleached to bright-pink in shells from beach drift. Pilsbry (1891, p. 93) was incorrect in as- suming that miniata is merely a rock-dwelling ecotype of the kelp living compressa. Description—Shell rather large, up to 93 mm. 3% inches) in length, strong but relatively thin, ovate, slightly attenuated in front, rather de- pressed, and with the apex varying between sub- A. W. B. Powell Patellidae central and the anterior third. Sculpture consist- ing of numerous primary radial cords, with mostly two radial threads in the interspaces; the ribbing varies in strength, and may be almost smooth to sharply-imbricated by dense concen- tric growth threads. Colour: externally radially streaked and speckled in reddish brown to bright- pink, on a white ground; internally silvery pinkish white, with the external pattern showing through strongly; spatula white-callused, sometimes tinged with orange. Living examples are usually en- crusted. Radula—Formula 3 + 1 + (2+1+2) + 1 +3. The radula stands nearest to that of compressa, the arrangement of the teeth being the same. The only noticeable difference between the two is in the shape of the cusps which have convex cutting edges in miniata but straight to concave ones in compressa. Plate 89. Patella (Cymbula) compressa Linnaeus, 1758. South Africa, 64-83 mm., AWBP coll. 1403 & 17985. [01-636] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 119 Plate 90. Figs. 1-3. Patella (Cymbula) miniata Born, 1778, South Africa. Figs. 1, 2. Port Nolloth, 75 mm., AWBP. coll. Measurements (mm.)— length width height 93.0 71.5 22.0 Natal; Mrs. N. Prior. 79.0 66.0 20.0 False Bay 75.0 56.5 16.0 Port Nolloth 54.0 43.5 12.0 Still Bay Synonymy 1778 Patella miniata Born, Index Mus. Caes. Vind., p. 436; 1780, Test. Mus. Caes. Vind., p. 420. 1786 Patella pulchra Lightfoot, Cat. Portland Mus., p. 105. 1791 Patella umbella Gmelin, Syst. Nat., ed. 13, p. 3706. 1791 Patella sanguinolenta Gmelin, Syst. Nat., ed. 13, p. 3716. 1798 Patella rosea Réding, Mus. Bolten., vol. 2, p. 9. 1798 Patella rubicunda Réding, Mus. Bolten, vol. 2, p. 9. 1848 Patella miniata Born, Krauss, Sudafr. Moll., Stuttgart, p. 51. 1854 Patella sanguinans Reeve, Conch. Iconica, pl. 6, fig. 10. 1854 Patella umbella Gmelin, Reeve, Conch. Iconica, pl. 9, figs. 17a, b. 1891 Patella. compressa var miniata Born, Pilsbry, Man. Conch., vol. 13, p. 94, pl. 26, figs. 22-27. 1932 Patella miniata Born, Turton, Mar. Shells Port Alfred, p. 168. 1932 Patella miniata umbella Gmelin, Turton, Mar. Shells Port Alfred, p. 168. 1932 Patella miniata decorata Turton, Mar. Shells Port Alfred, p. 169. pl. 39, fig. 1199. 1932 Patella pulchella Turton, Mar. Shells Port Alfred, p. 169, pl. 39, fig. 1200. (non Blainville, 1825). 1932 Patella alboguttata Turton, Mar. Shells Port Alfred, p. 169, pl. 39, fig. 1202. 1932 Patella denseplicata Turton, Mar. Shells Port Alfred, p. 169, pl. 39, fig. 1205. 1932 Patella densestriata Turton, Mar. Shells Port Alfred, p. 170, pl. 39, fig. 1206. 1933 Patella becki Turton, Journ. Conch., vol. 19, p. 371; nom. nov. pro P. pulchella Turton, 1932, non Blain- ville, 1825, 52455. Fig. 3. False Bay, 79 mm., AWBP coll. 26041. 1942 Patella sanguinolente (sic Gmelin, Tomlin & Stephenson, Proc. Malac. Soc., Lond., vol. 25, p. 7. 1949 Patella miniata Born, Koch, Ann. Natal Mus., vol. 11 (3), p. 506, pl. 21, figs. 1-12; text figs. 15a, b, 16 (radula). Records—SOUTH AFRICA: Port Nolloth in the west to Qolora in the east, common in the sub-littoral fringe (Koch, 1949, p. 506); Port Nolloth; False Bay; Still Bay; Port Elizabeth; Algoa Bay (all AWBP coll.); Natal (Mrs. N. Prior). Plate 91. Patella (Cymbula) miniata Born, 1778. Natal, South Africa; An extra large and fine example of the species, in the collection of Mrs. Nancy Prior of Cape Town. It has a length of 93 mm. (3% inches). [01-637] 120 Subgenus Cymbula A. W. B. Powell Patellidae [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence. | [01-635] November 27, 1973 Subgenus Olana H. and A. Adams, 1854 Type (monotypy): Patella cochlear Born, 1778 A moderate-sized shell of depressed pear- shape, with the anterior end laterally constricted and produced like a spout. The sole species of this subgenus is restricted to South Africa where in many places it is so abundant that it forms a dense mosaic, termed the “Cochlea zone.” A density of 1,300 examples to the square-yard has been recorded, and as many as 40 crowded on top of a single large shell. Almost all large examples bear several deeply excavated scars, resultant from superimposed individuals. These limpets do not appear to move around much, but merely rotate, so that the head can move in a circle and the radula crop the algal growth within its range (See Koch, 1949, pp. 498-499). Synonymy— 1854 Olana H. and A. Adams, The Genera of Recent Mol- lusca, vol. 1, p. 466. Type by monotypy: Patella coch- lear Gmelin, 1791 = Born, 1778. INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 121 Patella cochlear Born, 1778 (Pl. 63, figs. 12-14; Pls. 87, 92) Range—South Africa, from Buffalo River on the west coast and around the Cape of Good Hope to Port Edward, Natal. Description—Shell of moderately large size, up to 67 mm. (2% inches) in length, solid, depressed, pear-shaped, with the anterior end much con- stricted, and produced like a spout. Sculptured with strong, rather unequal radial ridges that deeply corrugate the margin. Colour: externally white to yellowish brown; internally white, tinged with bluish grey; the spatula fawn, orange-brown, or clouded with white, and surrounded, except in front of the head region, with a broad band of indigo. Radula—Formula 3 + 1 + (2+1+2) + 1 + 3. The small, slender, median central tooth, and the two pairs of multiple centrals form a horizontal line, as in true Patella, but the massive lateral has four cusps, two of them large and the other two much smaller. The cusps are heart-shaped, with a pronounced groove down the middle; marginals 3, each with a weak cusp. Measurements (mm.)—(all A. W. B. Powell col- lection). length width height 67.0 50.0 17.0 Port Alfred 58.5 43.5 15.5 Cape Natal 45.0 34.5 6.0 False Bay Plate 92. Patella (Olana) cochlea Born, 1778, Port Alfred, South Africa, 66 mm., AWBP coll. 51974. [01-643] 122. Subgenus Olana A. W. B. Powell Patellidae Synonymy— 1778 Patella cochlear Born, Index Mus. Caes. Vind., p. 437; 1780, Test. Mus. Caes. Vind., pl. 18, fig. 3, p. 420. 1790 Patella cochlear Born, Gmelin, Syst. Nat. ed. 13, p. 3721. 1848 Patella cochlear Born, Krauss, Sudafr. Moll., Stuttgart, p. 48. 1854 Patella cochlear Born, Reeve, Conch. Iconica, pl. 12, figs. 24a, b. [01-644] 1854 Patella (Olana) cochlear Born, H. & A. Adams, Gen. Rec. Moll., vol. 1, p. 466. 1891 Patella (Scutellastra) cochlear Born, Pilsbry, Man. Conch., vol. 13, p. 104, pl. 27, figs. 34, 35. 1949 Patella cochlear Born, Koch, Ann. Natal Mus., vol. 11 (3), p. 498, pl. 19, figs. 1-3; text figs. 5, 6 (radula). Records—SOUTH AFRICA: False Bay; Sea Point (Mrs. N. Prior); Port Elizabeth; Cape Natal; Port Alfred (all AWBP coll.). November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella = 123 nn. Subgenus Scutellastra H. and A. Adams, 1854 Type: Patella barbara Linnaeus, 1755 Mostly large massive shells with an opaque, non-iridescent, porcellaneous interior. The exter- nal coloration is either a uniform spread of colour, or the pigment may be confined to the spaces between the radial ribs and show through to the inside margin to form a narrow border where the shell is thinnest. Radula and gill cordon as in typical Patella, but the median central tooth is usually well-de- veloped, often as large as the other four centrals. The range of the subgenus is South Africa, on across the Indo-Pacific as far east as the Society Islands, northward to Japan and along the south coast of Australia. Undoubted fossil occurrences of the subgenus are cooperi (Powell, 1935) from the lower Miocene of Motuihi Island, Auckland, New Zealand, and aurorae Fleming, 1973, from the middle Oligocene of Mason River, North Canterbury, New Zealand. Synonymy— 1854 Scutellastra H. Adams and A. Adams, The Genera of Recent Mollusca, volume 1, p. 466, for gorgonica Hymphrey, pentagona Born and plicata Born. Type, by subsequent designation, Powell, 1938: Patella gor- gonica Humphrey “=P. longicosta Lamarck” (sic) =Pa- tella barbara Linnaeus, 1758. 1924 Patellanax Iredale, Proceedings of the Linnean Society of New South Wales, volume 49, part 3, p. 239. Type, by original designation: Patella squamifera Reeve, 1855. 1929 Penepatella Iredale, Memoirs of the Queensland Mu- seum, volume 9, part 3, p. 276. Type, by original designation: Penepatella inquisitor Iredale, 1929. Patella argenvillei Krauss, 1848 (PI. 65, fig. 4; Pls. 93, 96) Range—South Africa: Port Nolloth on the west coast, eastward to Qolora, between East London and Durban. Remarks—This species is more common on the west coast where it forms concentrated low-tidal communities, termed the Cochlear-argenvillei zone. It is one of the least variable of South African lim- pets, easily recognised by its narrowly elongate- oval, high-conical form, dense regular sculpture, and dark external coloration, with white linear rib interstices. Description—Shell large, up to 89 mm. (34 inches) in length, rather solid, oblong-ovate, slightly con- stricted at the anterior end, high conical, with the apex subcentral, a little nearer to the anterior end. Sculpture consisting of very numerous, over 100, more or less regular, flat-topped radial ribs, with linear interstices; margin delicately and evenly cren- ulated. Colour: externally blackish, the rib inter- stices white; internally white, dark-greyish between the marginal crenulations, and spatula diffused with yellowish brown. Radula—F ormula 3 + 1 + (2+1+2) + 1+ 3. Central teeth consisting of a narrow insignificant middle member, with a pair of strong, sharply-cusped and centrally-grooved teeth on each side; lateral massive, with an expanded top, bearing four sharp cusps, the middle two deeply grooved; marginals three, weakly cusped and slender. Measurements (mm.)— length width height 89.0 68.0 52.0 Table Bay rae) 58.0 35.5 South Africa 63.0 44.5 30.0 Port Elizabeth Plate 93. Patella (Scutellastra) argenvillet Krauss, 1545, Port Alfred, South Africa, 72-77 mim., AWBP coll. 30065. [01-649] 124. Subgenus Scutellastra Synonymy— 1848 Patella argenvillei Krauss, Sudafr. Moll., Stuttgart, p. 49; based upon Argenville Conch., 1870, vol. 1, p. 504 and vol. 3, pl. 3, fig. G. 1854 Patella argenvillii Krauss, Reeve, Conch. Iconica, pl. 10, figs. 20a, b. 1891 Patella (Scutellastra) argenvillei Krauss, Pilsbry, Man. Conch., vol. 13, p. 95, pl. 22, figs. 15, 16; pl. 58, fig. 44. 1949 Patella argenvillei Krauss, Koch, Ann. Natal Mus., vol. 11(3), p. 494, pl. 17, figs. 1-3; text figs. 1, 2 (radula). Records—SOUTH AFRICA: Table Bay (AWBP coll.); False Bay (AWBP coll.); Sea Point (Mrs. N. Prior); Port Elizabeth (Auck. Mus.); Port Alfred (AWBP coll.). Patella barbara Linnaeus, 1758 (Pl. 65, figs. 1-3; Pls. 94-96) Range—South Africa, the whole length of the coastline from Port Nolloth in the west to Umpan- gazi in the east. Remarks—This is a large, solid, ovate limpet, varying greatly in height, but always strongly radially ridged, resulting in a deeply corrugated margin. The coloration is buff to pale brownish externally and white within, except for the spatula, which is often blotched with reddish brown. Description—Shell rather large, up to 95 mm. (3% inches) in length, of only moderate height and weight in its younger stages but tall and crass in mature examples, narrowly to broadly ovate, with the apex near central. Sculpture variable, but always strongly and coarsely radially ribbed, their terminal points corrugating the margin, sometimes almost as Plate 94 Patella (Scutellastra) barbara Linnaeus, 1758, Port A. W. B. Powell Patellidae strongly as in longicosta. Radial ribs carinated and of varying strength, from 10 to 20 primaries and 1 to 4 secondaries in the interspaces, the whole rendered noticeably scabrous by close set, lamellose, concen- tric growth marks. The posterior end of the shell usually has 5 ribs much stronger than the rest. Colour: externally dull-buff to light yellowish brown; internally whitish, often with a narrow, pale-fawn, marginal border; spatula either irregu- larly blotched with reddish-brown, or callused over with white. A variable species, as shown by the lengthy synonymy. Radula—F ormula 3+ 1 + (2+1+2) + 1+3, similar to that of both longicosta and cochlear in the form of the massive lateral, which has an expanded head, bearing three cusps, the middle one much the larger, and there is an incipient fourth cusp on the outer side. The five centrals have the middle member small and slender, with a minute vestigial cusp, but the cusps of the outer pairs of centrals, and the middle member of the laterals have broad flat tops, with ridged margins; marginals 3, slender, flexuous, and each with a weak cusp. Measurements (mm.)— length width height 95.0 81.0 37.0 Buluga Bay, East London; Mrs. N. Prior 95.0 70.0 31.0 var. ovalis Pilsbry, 1891, p. 97 84.5 61.5 31.5 Still Bay 79.0 58.0 38.0 CC. of Good Hope 79.0 62.0 22.5 Port Alfred 72.0 60.0 27.0 Pilsbry, 1891, p. 96 Alfred, South Africa, 76 mm., AWBP coll. 30063. [01-650] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella = 125 Synonymy— 1758 Patella barbara Linnaeus, Syst. Nat., ed. 10, p. 782. 1778 Patella plicata Born, Mus. Caes. Vind., p. 433; 1780, Test. Mus. Caes. Vind., p. 417, pl. 18, fig. 1. 1786 Patella fungoides Lightfoot, Cat. Portland Mus., p. 55. 1786 Patella gorgonica Lightfoot, Cat. Portland Mus., p. 105. 1791 Patella plicaria Gmelin, Syst. Nat., ed. 13, p. 3708. 1791 Patella cypria Gmelin, Syst. Nat., ed. 13, p. 3698. 1819 Patella barbata Lamarck, Anim. sans vert., vol. 6 (1), p. 326. 1819 Patella spinifera Lamarck, Anim. sans vert., vol. 6 (1), p. 326. 1848 Patella barbara L., Krauss, Sudafr. Moll., Stuttgart, p. 45. 1848 Patellaa obtecta Krauss, Sudafr. Moll., Stuttgart, p. 47, pl..3, fig. 11. 1854 a le Born, Reeve, Conch. Iconica, pl. 9, figs. l6a, b. 1891 Patella (Scutellastra) barbara L., Pilsbry, Man. Conch., vol. 13, p. 96, pl. 15, figs. 1, 2; pl. 59, figs. 50-55. 1591 Patella (Scutellastra) barbara var. ovalis Pilsbry, Man. Conch., vol. 13, p. 97, pl. 60, figs. 56-58. 1932 Patella barbara L., Turton, Mar. Shells Port Alfred, p. 163, sp. 1162. 1932 Patella barbara plicata Born, Turton, Mar. Shells Port Al- fred, p. 163. 1932 Patella whitechurchi Turton, Mar. Shells Port Alfred, p. 164, pl. 37, fig. 1165. 1932 Patella thetis Turton, Mar. Shells Port Alfred, p. 164, pl. 37, fig. 1166. 1932 Patella hera Turton, Mar. Shells Port Alfred, p. 164, pl. 37, fig. 1167. 1932 Patella amphitrite Turton, Mar. Shells Port Alfred, p. 164, pl. 37, fig. 1168. 1932 Patella amphitrite var. brunescens Turton, Mar. Shells Port Alfred, p. 164, pl. 37, fig. 1169. 1932 Patella decemcostata var. major Turton, Mar. Shells Port Alfred, p. 165, pl. 35, fig. 1171. 1932 Patella nympha Turton, Mar. Shells Port Alfred, p. 165, pl. 38, fig. 1174. 1932 Patella sowerbyi Turton, Mar. Shells Port Alfred, p. 166, pl. 38, fig. 1176. 1949 Patella barbara Linne, Koch, Ann. Natal Mus., vol. 11 (3), p. 496, pl. 18, figs. 1-12; text figs. 3a, b, 4 (radula). Records SOUTH AFRICA: whole length of coastline from west to east (Koch, 1949, p. 496); Table Bay (Auck. Mus. 3081); Cape of Good Hope; Still Bay; False Bay; Port Al- fred; Cape Natal (all AWBP coll.). Patella longicosta Lamarck, 1819 (Pl. 65, figs. 5-7; Pls. 95, 96) Range—South Africa, from Oudekraal, west side of Cape Peninsula, eastward to Umpangazi, north of Durban. Remarks—This species is easily recognised by its depressed star-shape, with the primary ribs extend- ing well-beyond the margin, which is narrowly bor- dered in black. This species occurs commonly in the lower Balanoid and Cochlear zones. Description—Shell rather large, up to 76 mm. (3 inches) in length, solid, depressed, stellate, very strongly sculptured with sharply carinated, radial ridges, that project well-beyond the margin, seven of them much stronger than the rest; apex at an- terior third to submedian. Colour of exterior dull- black, when not eroded, to a rusty-brown; interior bluish white, with a narrow black margin, and a yellowish brown spatula, the latter clouded with a white callus in senile specimens. Radula—F ormula 3+ 1 + (2+1+2) + 1+3, somewhat similar to that of cochlear in the form of the cusps, which are leaf-shaped with a median groove, and in the massive lateral that has four cusps. Plate 95. Patella (Scutellastra) longicosta) Lamarck, 1519. Mossel Bay, Cape Peninsula, South Africa, 66 mm., AWBP coll. 42429. [01-651] 126 Subgenus Scutellastra A. W. B. Powell Patellidae a Se ee Measurements (mm.)— 1901 Patella decemcostata E. A. Smith, Journ. Conch., vol. 10, . p. 106, pl. 1, fig. 22. length width height 1932 Patella longicosta Lam., Turton, Mar. Shells Port Alfred, 75.5 75.5 23.0 False Bay p. 161, pl. 37, fig. 1145. 70.0 64.0 18.0 False Bay 1932 Patella longicosta intermedia Turton, Mar. Shells Port Al- 61.0 56.0 12.0 Cape Peninsula fred, p. 161, pl. 37, fig. 1146. 1932 Patella multilirata Turton, Mar. Shells Port Alfred, p. 161, Synonymy— pl. 37, fig. 1147. 1932 Patella tabularis Krauss, Turton, Mar. Shells Port Alfred, p. 1819 Patella longicosta Lamarck, Anim. sans vert., vol. 6 (1), 161. (non Krauss, 1848; in part, smaller of two examples). p. 326. 1932 Patella tabularis angulosa Gmelin, Turton, Mar. Shells 1842 Patella longicosta Lam., Reeve, Conch. Syst., vol. 2, p. 15, Port Alfred, p. 161, sp. 1150. (non Gmelin, 1791). pl. 136, fig. 6. 1932 Patella tabularis monopsis Gmelin, Turton, Mar. Shells Port 1848 Patella longicosta Lam., Krauss, Sudafr. Moll., Stuttgart, Alfred, p. 162. (sic; non monopis Gmelin, 1792). p. 48. 1932 Patella tabularis squamosa Gmelin, Turton, Mar. Shells 1854 Patella longicosta Lam., Reeve, Conch. Iconica, pl. 6, figs. Port Alfred, p. 161, sp. 1149. (non Gmelin, 1791). lla, b. 1932 Patella granatina Linn. Turton, Mar. Shells Port Alfred, 1891 Patella (Scutellastra) longicosta Lamarck, Pilsbry, Man. p. 163, sp. 1161. (non Linnaeus, 1758). Conch., vol. 13, p. 107, pl. 28, figs. 37, 38. 1932 Patella decemcostata E. A. Smith, Turton, Mar. Shells Port Alfred, p. 165. 1932 Patella albanyana Turton, Mar. Shells Port Alfred, p. 165, pl. 38, fig. 1175. 1933 Patella longicosta kowiensis Turton, Journ. Conch., vol. 19, p. 371; nom. nov. pro P. longicosta intermedia Turton, 1932, non Knapp, 1857. 1942 Patella longicosta Lam., Tomlin and Stephenson, Proc. Malac. Soc., Lond., vol. 25, pp. 4-9. 1949 Patella longicosta Lam., Koch, Ann. Natal Mus., vol. 11 (3), p. 504, pl. 20, figs. 6-13; text figs. 13a-c, 14 (radula). Records—SOUTH AFRICA: Cape Peninsula (AWBP coll.); Kommetje, Cape Peninsula (Mrs. N. Prior); Mossel Bay, Cape Peninsula (V. Orr, 1955; ANSP); False Bay (AWBP coll.); Still Bay (Auck. Mus.); Simon’s Bay (V. Orr, 1955; ANSP); Port Eliza- beth (Auck. Mus.); Cape Natal (AWBP coll.); Port Alfred (AWBP coll.). Types—The type series of three examples of longi- costa is in the Muséum D’Histoire Naturelle de Geneve. Patella tabularis Krauss, 1848 (Pl. 66, fig. 5; Pls. 96, 97) Range—South Africa, from Cape Peninsula east- ward to Port St. John’s. Remarks—This is the largest of the South African limpets; it somewhat resembles kermadecensis but has much more prominent radial sculpture. The species inhabits the sub-littoral fringe, and does not occur in dense communities (Koch, 1949, p. 509). Description—Shell very large and massive, up to 147.5 mm. (5% inches) in length, broadly ovate, mod- erately elevated, and with the apex anterior to the middle. Sculpture heavy and coarse, consisting of 9, 10 or more heavy, foldlike, radial ribs, and nu- merous secondary ribs of several sizes, the whole imbricated by dense lamellose growth lines; margin deeply and somewhat irregularly scalloped. Colour: exterior dull reddish or rusty-brown; interior porcel- Plate 96. Fig. 1. Radula of Patella (Scutellastra) barbara lanous-white, with a moderately wide border of Linnaeus. Fig. 2. Patella (Scutellastra) longicosta Lamarck. ans : . ; ‘ = Fig. 3. Patella (Seutellastra) argenvillei Krauss. Fig. 4. reddish brown, being the external colour show ane Patella (Scutellastra) tabularis Krauss. South Africa. Radulae, through at the thinner margin; spatula not differen- all from: Koch, 1949. tiated by colour. [01-652] November 27, 1973 Plate 97. Patella (Scutellastra) tabularis Krauss, 1848. Port Alfred, South Africa, 115 mm., AWBP coll. 30062. Measurements (mm.)— length width height 147.5 125.5 50.0 Buluga Bay, East London; Mrs. N. Prior 127.0 111.9 46.0 South Africa 114.0 98.0 32.0 Cape Point 81.0 68.0 20.5. Port Alfred Radula—F ormula 3 + 1 + (2+1+2) + 1 + 3. Central teeth consisting of a slender, almost vestigial, middle member, flanked by pairs of large, blunt-cusped INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella = 127 teeth; lateral massive, with an expanded head, bear- ing four cusps, the middle pair the stronger; mar- ginals three, narrow and slender, each with a weak cusp. Synonymy— 1848 Patella tabularis Krauss, Sudafr. Moll., Stuttgart, p. 47, pl: 3; fig. 8. 1854 Patella rustica L., Reeve, Conch. Iconica, pl. 5, figs. 8a, b. (non Linnaeus, 1758). 1891 Patella (Scutellastra) tabularis Krauss, Pilsbry, Man. Conch., vol. 13, p. 105, pl. 16, figs. 9, 10. 1891 Patella (Scutellastra) patriarcha Pilsbry, Man. Conch., vol. 13, p. 105, pl. 64, figs. 84, 85; pl. 65, fig. 86. 1932 Patella rustica L. Turton, Mar. Shells Port Alfred, p. 162, sp. 1154. 1932 Patella tabularis Krauss, Turton, Mar. Shells Port Alfred, p. 161, sp. 1148 (in part; larger of two specimens). 1932 Patella obtecta Krauss Turton, Mar. Shells Port Alfred, p. 162, sp. 1155. (non Krauss). 1932 Patella patriarcha Pilsbry, Turton, Mar. Shells Port Alfred, p. 162, sp. 1153. 1942 Patella tabularis Krauss Tomlin and Stephenson, Proc. Malac. Soc., Lond. vol. 25 (1), p. 5. 1949 Patella tabularis Krauss, Koch, Ann. Natal Mus., vol. 11 (3), p. 509, pl. 22, figs. 5-11; text figs. 19a, b, 20 (radula). Types—The type of tabularis is in the Stuttgart Mu- seum, formerly the Naturalien Cabinet. Records—SOUTH AFRICA: Cape Peninsula to Port St. John’s (Koch, 1949, p. 509); Cape Point (AWBP coll.); Port Alfred (AWBP coll.); Buluga Bay, East London (Mrs. N. Prior). Patella exusta Reeve, 1854 (Pl. 9S, tig. 1; Pl. 104, fig. 3) Range— Mauritius. Remarks—The typical subspecies seems to be confined to Mauritius, although the wider-rang- ing subspecies pica Reeve, sometimes occurs along with it. The distinctive character of exusta is the dull-black, thick outer layer of the shell, which also produces the narrow, black internal border. It is unfortunate that the better known name, chitonoides has to fall as a synonym of exusta. Description—Shell of moderate size, up to54 mm. (2% inches) in length, solid, rather depressed, elon- gate-ovate, and decidedly narrowed anteriorly. Apex subcentral to about the anterior third. Sculpture consisting of about 20 narrow sharply raised primary radial ribs, and 3 or 4 weak radial threads in the wider interspaces, over the post- erior half of the shell. The margin is broadly and shallowly corrugated. Colour of exterior, when not encrusted or abraded, uniformly dull black; interior pale bluish to pinkish white, with a nar- row black margin. The spatula is weakly defined, and sometimes partly clouded by a pale fawn callus. [01-653] 128 Subgenus Scutellastra Measurements (mm. )— length width height 54.0 38.0 13.0 syntype of chitonoides 53.0 44.0 14.0 lectotype of exusta 48.0) 38.0 16.0 Mauritius 47.0 37.75 11.5. lectotype of chitonoides 45.0 35.0 12.5. Mauritius Synonymy— 1854 Patella exusta Reeve, Conch. Iconica, pl. 15, figs. 35a, b. (locality unknown). 1854 Patella chitonoides Reeve, Conch. Iconica, pl. 21, figs. 52a, b. (locality unknown). Dec. 1554. 1891 Patella (Scutellastra) exusta Reeve, Pilsbry, Man. Conch. vol. 13, p. 98, pl. 24, figs. 9, 10. 1891 Patella (Scutellastra) pica form chitonoides Reeve, Pilsbry, Man. Conch., vol. 13, p. 98, pl. 26, figs. 28, 29. Types—The types of both exusta and of chito- noides are in the British Museum (Natural History). That of exusta consists of three syntypes glued to a tablet. One of these, measuring 53 x 44 x 14 mm., matches Reeve’s figure, pl. 15, fig. 35, and is here nominated lectotype. The other two specimens on the tablet are the Californian Acmaea pelta Esch- scholtz, and may have been added later than Reeve. The type series of chitonoides also consists of three syntypes, and the one matching Reeve’s fig. 52, Plate 98. Fig. 1. Patella (Scutellastra) exusta Reeve, 1554. Mauritius, 48 mm., AWBP. coll. 628. Figs. 2-4 Patella 9 (Scutellastra) exusta subspecies pica Reeve, 1854. Fig. 2. Mauritius, 38 mm., AWBP. coll. 627. Figs. 3.4. Mahé, Seychelles, 42-45 min., AWBP coll. 46135. A. W. B. Powell Patellidae measuring 47 x 37.75 x 11.5 mm. is here nomi- nated lectotype. Records—Locality unknown for the types of both exusta and chitonoides. MAURITIUS: (AWBP coll.); Vacoas Point, 3 miles south of Mahebourg (Ruth Ostheimer and Virginia Orr, Nov. 20, 1960). Patella exusta subspecies pica Reeve, 1854 (Pl. 98, figs. 2-4) Range—Islands of the Indian Ocean, Mauritius to Seychelles. Description—Shell of similar size and shape to exusta typical, being elongated and noticeably nar- rowed anteriorly, but the coloration is different, for instead of the whole of the outer surface being black, that colour is confined to radiate lines or streaks, often short and intermittent, and present only in the primary rib interstices, the rest of the exterior of the shell being white. Internally the shell is porcellanous-white with the spatula irregularly stained orange-brown, and at the margin the dark external pattern shows through, forming an inter- mittent narrow border. Measurements (mm.)— length width height 49.5 38.00 14.0 — lectotype 47.0 37.25 10.0 Mauritius 44.5 36.25 10.5 Mahe, Seychelles 38.0 30.00 6.5. Mauritius 30.4 26.00 5.0 Seychelles Synonymy— 1854 Patella pica Reeve, Conch. Iconica, pl. 19, figs. 45a-c. 1891 Patella (Scutellastra) pica Reeve, Pilsbry, Man. Conch., vol. 13, p. 97, pl. 22, figs. 9, 10, 13, 14; pl. 59, figs. 47- 49 (not pl. 26, figs. 28, 29). 1863 Patella moreli Deshayes, Cat. Moll. Réunion, p. 43, pl. 6, fig. 13. 1863 Patella levata Deshayes, Cat. Moll. Réunion, p. 44, pl. 6, fig. 14. Records—*‘South Seas” (type); MAURITIUS: (AWBP coll.); Vacoas Point, 3 miles south of Mahebourg (Ruth Ostheimer and Virginia Orr, Nov. 20, 1960). SEYCHELLES: (AWBP coll.); Mahé (AWBP coll.); Frigate Island (AWBP coll.). REUNION: (Deshayes, 1863). Types—The type series of pica is in the British Museum (Natural History) and consists of three syn- types mounted upon a tablet. The one measuring 49.5 x 38 x 14 mm. is here nominated lectotype. [01-654] November 27, 1973 Patella flexuosa Quoy and Gaimard, 1834 (Pl. 65, figs. 8, 9; Pls. 99, 104) Range—Andaman Islands and the tropical Pa- cific as far east as the Tuamotu Islands. Remarks—This exceedingly variable species has had many names, and unfortunately the one under which it has usually appeared, stellaeformis Reeve, 1842, must fall as a synonym of the earlier flexuosa Quoy and Gaimard, 1534. The species is intertidal and a shallow-water dweller, living attached to coral rock, or some- times upon the outer and inner surfaces of large shells. Examples living attached to shells tend to be of lighter build, and of more circular outline INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella = 129 than those from coral-rock faces, but that is not invariably so, and all manner of shapes, sculpture and coloration is encountered, irrespective of station. The species varies between 14 mm. and 41 mm., is circular to elongate-ovate in shape, may be almost flat to moderately elevated, thin or solid. The strength of the external ribbing deter- mines the degree to which the margin is scal- loped. The exterior is dull-white, sometimes spar- ingly speckled, lined, or with zigzag dark-brown markings in the rib interstices, but the outer surface is usually lime-encrusted. The interior is porcellanous, more or less white, and the spatula Plate 99. Figs. 1-9. Patella (Scutellastra) flexuosa Quoy & Gaimard, 1834. Fig. 1. Rouw Island, Aoeri Islands, West New Guinea (stellaeformis form), 30 mm., AWBP. coll. 39227. Figs. 2, 3. Michaelmas Cay, off Cairns, North Queensland, 29-34 mm. (topotypes of inquisitor), AWBP coll. 46062. Fig. 4. New Caledonia, 31 mm. (topotype of intraurea), AWBP. coll. 45672. Figs. 5, 6. Wake Island, 31-34 mm., AWBP. coll. 204725. Figs. 7, 8. Paea, Tahiti, Society Islands 33-35 mm. (paumotensis form), AWBP coll. 115. Fig. 9. Bikini Island, Marshall Islands, 35 min., AWBP coll. 48226 [01-655] 130. Subgenus Scutellastra A. W may be white, yellowish, orange-brown, or some- times dark-slate. Description—Shell rather small, up to 42 mm. (1% inches) in length, typically roundly-octagonal, with the apex nearly central, sculptured with 8 or 9 rounded radial folds that project strongly at the margin. The entire surface, folds and interstices alike, is densely sculptured with secondary, crisp, scabrous to spinose cords. In some forms the pri- mary and secondary radials become nearly equal, in which cases the octagonal outline is modified to a crenulated oval. Colour as described under remarks. Radula—Formula 3 + ] + (2+1+2) + 1 + 3, very similar to that of exusta pica and kermadecensis. Prashad and Rao figured the radula of their ‘tara, which compares closely with the radula of a Cook Islands specimen. Prashad and Rao con- sidered their species to be related to the South African granularis, but in that species the cusps are better formed, parrot-beaklike, and the shell is quite unlike the flexuosa group in form, size, texture and coloration. Measurements (mm.)— length width height 41.0 35.0 17.5 Fakarawa, Tuamotus 39.0 30.0 9.5 ‘Tokorava, Tuamotus 33.0 30.0 8.0 Michaelmas Cay (type of inquisitor) 30.0 28.0 9.0 HICOBARL 4.) 8. 3 CEYLON | : *. Mojuro CAROLINE is! | ‘s MALDIVE | Makin | - y FLEXUOSA FLEXUOSA ce o° = — —— oo ee TF ty | So, Pa ' - CHAGOS Veo, aa | PHOENIX 15. | . _At 5 “Qs “Ce TOKELAU IS _MARQUESAS IS. comonse EXUSTA PICA €0C06 - KEELING. os = . gr rots + : SAMO ig” ? : I: es y CH, NEW SB VHEBRIE al e : e& R . 2 2 So, or Gare a | é S05: hs FISTS. B fo Cre 20° ——— oe = + 2 OS = — aa ———— | i ne ¥ Filan. 4 | ea bee cat ee | ® 2 __ ge: ae J See | | . | on J | Putcairn | Norkolk | i fous | ‘| | H | | NEW t | ZEALAND Mania Theresa n _ n 1 = atin a 40 0 80 100 160 180° i) 140° Plate 101. Geographical distribution of Patella (Scutellastra) exusta Reeve, Patella (Scutellastra) exusta subspecies pica Reeve, Patella (Scutellastra) flexuosa Quoy & Gaimard, and Petella (Scutellastra) flexuosa subspecies optima Pilsbry. [01-655] 132. Subgenus Scutellastra Records—JAPAN: Yakushima, Osumi (type); Suwanose- jima, Osumi (AWBP coll.); Waki, Kyushu Island (AWBP coll.). Amami and Ryukyu Islands (Habe, 1964). Types—The holotype is in the Academy of Natural Sciences of Philadelphia. No. 98023. Patella kermadecensis Pilsbry, 1894 (Pl. 66, fig. 1; Pls. 102-104) Range—Kermadec Islands Remarks—This species is restricted to the Ker- madec group, and is the second largest known living member of the Patellidae, sometimes at- taining a length of seven inches, and exceeded in size only by the tropical West American Patella (Ancistromesus) mexicana. The white porcellanous interior, and orange border of the shell, as well as its broadly ovate shape, readily distinguish kermadecensis. Description—Shell very large and massive, up to 174 mm. (6% inches) in length, broadly ovate, but slightly narrowed in front, apex almost central, and dorsal slopes almost straight. Sculpture con- sisting of a dense coverage of narrow radial ribs, all rendered weakly scabrous by concentric growth lines. The primary radials number from 30 to 50, and the secondary ones, which are al- most as strong, number from 3 to 5 for each inter- space. The margin is broadly and_ shallowly scalloped, corresponding to the interspaces of the primary radials. Colour of exterior dull-orange; interior porcellanous-white, with a narrow mar- gin, that varies from pale to bright orange, and very occasionally is dark greenish-brown. The muscle impression is mostly dull cream, but occasionally it is greenish. Young shells have the A. W. B. Powell Patellidae spatula fawn to pale orange-brown, and the or- ange border to the shell, varying from plain or- ange or radially streaked to almost continuously blotched with brown. The greenish muscle im- pression is present only in shells that have been thinned by external erosion. Radula—Formula 3 + 1 + (2+1+2) + 1 +3. Radula very short and folded back upon itself at the end, as in typical Patella. There are five central teeth in a horizontal row, the median one_ shorter, narrower, and much smaller than the other four; all five bear blunt chisel-shaped cusps. The lateral is massive, with a broad head, bearing three fused chisel-shaped cusps. The three semitransparent marginals are small, elongated and narrow, each with a small blunt cusp. Animal—As in Patella vulgata, the gill cordon is continuous, not interrupted by the head as in Cellana, and the cephalic tentacles are short and broadly conical, with the eye in a pit at the outer base. Measurements (mm.)— length width height 174.0 160.0 — Oliver, 1915, p. 510 153.0 137.0 51.0 Raoul Island 148.5 135.0 50.0 Reoul Island 136.0 117.0 42.0 Raoul Island 123.5 108.0 30.5 Raoul Island Synonymy— 1894 Patella (Scutellastra) kermadecensis Pilsbry, The Nautilus, vol. 7, p. 109. 1894 Patella kermadecensis Pilsbry, Taylor, The Nautilus, vol. 7, p. 142. 1894 Patella pilsbryi Brazier, Proc. Linn. Soc. N. S. W., vol. 9, ser. 2, p. 183 (disputed locality, substituted South Africa, and renamed species). Plate 102. Patella (Scutellastra) kermadecensis Pilsbry, 1894. Raoul Island, Kermadec Islands, 123.5 mm., AWBP coll. 207. [01-656] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella = 133 1894 Patella kermadecensis Pilsbry, Proc. Acad. Nat. Sci. Phila. pp. 208-212, pls. 7, 8 (Kermadec locality con- firmed). 1895 Patella (Scutellastra) kermadecensis Pilsbry, Cheese- man, Proc. Linn. Soc. N. S. W., vol. 10, ser. 2, pp. 221-223 (Kermadec locality confirmed). 1902 Patella kermadecensis Pilsbry, Suter, Journ. Malac., vol. 9, p. 111, pl. 8 (animal and dentition). 1915 Scutellastra kermadecensis Pilsbry, Oliver, N. Z. Inst., vol. 47, p. 510. Types—The type specimens are in the Academy of Natural Sciences of Philadelphia. Records —KERMADEC ISLANDS: Raoul (Sunday Island) (type); Raoul, Macaulay, and Curtis Islands, also French Rock (Oliver, 1915); Raoul Island (Auck. Mus.); (AWBP coll.); north and east coasts of Raoul Island, also Meyer Islet (Cheese- man, 1895). Trans. Patella aurorae Fleming, 1973 (PI. 105) Range—New Zealand; Mason River, north of Waiau, North Canterbury, in a boulder derived from the Isolated Hill Limestone of Duntroonian Oligocene age. Remarks—This large and massive New Zealand fossil limpet is an obvious forerunner of the Recent kermadecensis, now confined to the Kermadecs, which in turn is related to the Melanesian tuco- piana. These isolated occurrences, both in time and in location, suggest that this group of limpets once had a considerable geographical range. Plate 103. Figs. 1-5. Patella (Scutellastra) kermadecensis Pilsbry, 1894, Raoul Island, Kermadec Islands. Young stages in ascending order of size, from Fig. 1, length 30 mm. to Fig. 5, length 67 mm. Figs. 2 and 5 have a plain orange border. Figs. 1, 3 and 4 have the orange border streaked with dark-brown. Figs. 3 and 4 are elevated, but Fig. 5 is very depressed. AWBP coll. 207, 17764 and 25116. The Oligocene fossil, compared with the Recent kermadecensis is less noticeably narrowed in front, has the apex nearer to the anterior margin, and the sculpture is very much stronger. These differences are here considered to war- rant full specific, rather than subspecific status for aurorae. Description—Shell very large and massive, up to 200 mm. (5 inches) in length, broadly ovate and moderately elevated, height a little less than one third that of the length. Sculpture coarse, consisting of about 32 strong primary radials and 4-6 secondary radials in each interspace. Apex about two fifths the length, from the anterior end, which is only slightly narrowed. Measurements (mm. )— length width height 180 150 78 holotype 200 — paratype Types—Holotype and paratype in the collection of the New Zealand Geological Survey, Lower Hutt, Wellington. Synonymy— 1973 Patella (Scutellastra) kermadecensis aurorae’ Fleming, N.Z. Journ. Mar. & Freshw. Res. vol. 7 (1 & 2), p. 160. Plate 104. Fig. U1. Patella (Scutellastra) kermadecensis Pils- bry, Raoul Island, Kermadec Islands. Radula. Fig. 2. Patella (Scutellastra) flexuosa Quoy & Gaimard, Rarotonga, Cook Islands. Radula. Fig. 3. Patella (Scutellastra) exusta sub- species pica Reeve, Mauritius. Radula. (01-657 | 134. Subgenus Scutellastra A. W. B. Powell Patellidae 2 sg Ag Ss of oe fis Plate 105. Patella (Scutellastra) aurorae’ Fleming, 1973. Mason River, North Canterbury, New Zealand. Duntroonian Pligocene. Patella tucopiana (Powell, 1925) (Pl. 106) Range—Tikopia (Tucopia) Island, Melanesia. Remarks—A large solid limpet, but evidently of smaller adult size than kermadecensis, from which it differs in shape, being regularly ovate, not narrowed anteriorly, and also in the coloration of the exterior, which is black instead of orange. The writer knows of only the two type specimens of this rare species. They were obtained from a native of Tikopia who stated that they came from a nearby reef. Description—Shell large, up to 92 mm. (3% inches) in length, solid, ovate, depressed, the apex at about the anterior third, anterior slope almost straight, posterior slope convex, margin weakly crenulated. Sculpture consisting of num- erous low narrow irregular radial ribs, 10 of them primary, and between 80 and 85 secondary radials. Colour of exterior dull black, more deeply impregnated in the rib interstices; interior creamy- white, with the spatula tinged pale flesh-colour; margin with a narrow amber coloured border, through which the external colour shows as a continuous series of irregular black dashes. Measurements (mm.)— length width height 92.0 73.0 25.0 holotype $1.0 63.0 22.0 paratype [01-658] Synonymy— 1925 Scutellastra tucopiana Powell, Proc. Malac. Soc., London, vol. 16, pt. 4, p. 169. Types—The holotype and paratype are in the Powell collection, Auckland. Known only from the type locality. Plate 106. Patella (Scutellastra) tucopiana (Powell, 1925). Vikopia Island, Melanesia. Holotype (above), 92 mm., and paratype, 81 mm., AWBP coll. 206. November 27, 1973 Plate 107. Figs. 1-4. Patella (Scutellastra) laticostata Blain- ville, 1825. Figs. 1, 2. Yellengap, Western Australia, 46-50 mm., AWBP coll. 51976. Figs. 3, 4. Cape Naturaliste, Western Australia, $1.5 mm., AWBP coll. 29117. Patella laticostata Blainville, 1825 (Pl. 66, figs. 3, 4: Pls. 107 and 113) Range—Western South Australia to southern Western Australia; lower littoral zone. Remarks—This is the largest member of the subgenus Scutellastra found in Australian waters. INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella = 135 It is easily recognised by its solidity, large size, often over four inches in length, narrowly-ovate shape and high-conical profile. The interior is creamy-white, varyingly stained with orange- brown, and there is a marginal border of dark- brown maculations. The exterior is nearly always eroded, but in fresh non-eroded examples there is a coarse radial sculpture of dark-brown ribs. Description—Shell very large, up to 110 mm. (4% inches) in length, very solid, narrowly-ovate, and frequently high in profile, the apex at about the anterior third. Radial ribbing coarse and irregular in size; juveniles with about 22 primary ribs, in- creasing by interpolation to about 50 primaries in the adult, and there are finer subsidiary radials in the interstices. Interior porcellanous, with the crenulated border variably maculated, and with a well-defined, very large spatula, mainly white, or diffused with yellowish brown, but often sur- rounded at its outer edge by an irregular zone of deep orange-brown. The margin is rather wide in young shells but relatively narrow in the fully adult, and bears numerous radiate lines or thick dashes, in dark-brown, on a whitish ground. Ex- ternally the shell is usually eroded to a dull green- ish grey, and sometimes bears one or two speci- mens of the acmaeid, Patelloida nigrosulcata (Reeve), which deeply excavate the surface. Radula—Formula 3 + 1 + (2+1+2) + 1 + 3. There are five central teeth, the middle one as long as, but narrower than, the other four, each with a single well-developed shovel-shaped cusp; the large palmate lateral bears four blunt cusps, and each of the three, slender, rod-like marginals has a blunt poorly developed cusp (see Macpherson, 1955, p. 235). 120° 140° Plate 10S. Geographical distribution of Patella (Scutellastra) laticostata” Blainville, Patella (Scutellastra) kermadecensis Pilsbry, Patella (Scutellastra) tucopiana (Powell), and the Br | . a L Ss : ee | gv 3 " PHOENIX 18 Re. “Ce ", TOKELAU 1S > 3s Tikopia ‘S 5, . Ss z : \ TUCOPTANA | “204 15 , oy NEW ars : ; ° . . a S, . f [e) fe Ce TONGA = cook |. * Sea Sa is.t_— -“4T<- | Usy, | KERMADEC Is. | KERMADECENSIS COOPERI Miocene 180° New Zealand Miocene Patella (Scutellastra) cooperi (Powell) These are all large species, Comparable in size with the largest of the South African members. [01-659] 136 = Subgenus Scutellastra Measurements (mm.)— length width height 110.0 85.0 54.0 all Charley Island, 92.0 68.0 35.0 Archipelago of the 83.0 60.0 56.0 Recherche 77.0 53.5 29.0 Synonymy— 1825 Patella laticostata Blainville, Dict. Sci. Nat., vol. 38, p. 111. 1826 Patella neglecta Gray, King’s Intertropical Survey Aust., vol. 2, pp. 156, 182, 492. 1843 Patella rustica Menke, Moll. New Holl., p. 33. 1854 Patella zebra Reeve, Conch. Iconica, vol. 8, pl. 4, figs. 7a, b. Swan River. (non Blainville) 1891 Patella (Scutellastra) neglecta’ Gray, Pilsbry, Man. Conch., vol. 13, p. 95, pl. 20, fig. 41. 1924 Patella laticostata Blainville, Iredale, Proc. Linn. Soc. N.S.W., vol. 49, p. 241. 1955 Patellanax laticostata Blainville, Macpherson, Proc. Roy. Soc. Vict., vol. 67 (2), p. 234, text figs., pl. 8, fig. 4. 1959 Cellana laticostata Blainville, Cotton, S. Aust. Moll., Archaeogast., Govt. Print., Adelaide, p. 292. Records—Western SOUTH AUSTRALIA: Port Lincoln and Streaky Bay (B. C. Cotton, 1959). South WESTERN AUS- TRALIA: King George Sound (B. C. Cotton, 1959); Charley Island, Archipelago of the Recherche; Quarantine Ground, Albany; Foul Bay; Cape Naturaliste; Garden Island, Fre- mantle (all AWBP coll.). A record from Shark Bay, North Western Australia (B. C. Cotton, 1959) requires confirmation. Patella peronii Blainville, 1825 (Pl. 65, figs. 11-13; Pls. 109 and 113) Range—Australia; from southern Western Aus- tralia to Victoria, Tasmania and New South Wales. Remarks—This species, also, is extremely var- iable in shape and in sculptural development, ranging from typical peronii, which is weakly but regularly smooth ribbed, thus producing an al- most smooth margin, to the strongly squamose sculptured squamifera form, in which the margin becomes noticeably corrugated. Dr. Hope Mac- pherson (1955) claimed that when large series were examined, both forms were seen to inter- grade, and no differences in the radula were ap- parent. Common on exposed rock platforms of the lower littoral zone to the sublittoral fringe, among holdfasts of giant kelp. Description—Shell of moderate size, averaging about 35 mm. but reaching 47 mm. (1% to Ik inches) in length, solid, tall-conical, with the apex varying between subcentral and the anterior fourth. Sculpture extremely variable, ranging from low, rounded, primary radials with 4 to 6 radial threads in the interspaces (typical form), to A. W. B. Powell Patellidae strongly sculptured shells with about 24 bold carinated and spinose primary radials, with a few relatively strong interstitial radials (forma squami- fera). Colour, externally yellowish brown with the radials paler, interior porcellanous-white with a faint yellowish brown spatula. In the typical peronii form the secondary radial interspaces are often lined in black. Shell margin thin semitrans- parent, yellowish with the external interstitial lines showing through. The shell margin varies according to the strength of the external sculpture being almost smooth in the peronii form but strongly corrugated in the squamifera form. Radula—Formula 3 + 1 + (2+1+2) + 1+ 3. Central teeth five, the middle one much smaller than the other four, lateral with four cusps, followed by the usual three functionless marginals (Macpher- son, 1955, p. 233). Measurements (mm.)—(A) = typical peronii; (B) = squamifera form. length width height 47.0 39.0 24.0 Port Arthur, Tasmania; (A) 43.5 34.0 16.0 — Port Jackson; (B) 37.4 29.5 15.0 New South Wales; (B) 31.5 25.0 12.5. Shellharbour, N.S.W.; (B) Plate 109. Figs. 1-4. Patella (Scutellastra) peronii Blainville, 1825. Figs. 1, 2. Merimbula, New South Wales, 39-42 mm., AWBP. coll. 51071. Fig. 3. Port Jackson, New South Wales, 43 mm., AWBP. coll. 212 (squamifera form). Fig. 4. Port Arthur, Tasmania, 47 mm., AWBP coll. 45421 (peronii = ustulata form). [01-660] November 27, 1973 ue NEW Dvy : | °° EBRIDES 4 , « . . * ou eS new e Ca CALEDONIA | _ FANS. Plate 110. Geographical distribution of Patella (Scutellastra) peronii Blainville. Synonymy— 1825 Patella peronii Blainville, Dict. Sci. Nat., vol. 38, p. 111. 1848 Patella diemenensis Philippi, Zeitsch. f. Malak., vol. 5, p. 162. 1855 Patella ustulata) Reeve, Conch. Iconica, pl. 31, figs. $8a, b. 1855 Patella aculeata Reeve, Conch. Iconica, pl. 32, fig. 90 (non Gmelin). 1855 Patella squamifera Reeve, Conch. Iconica, pl. 32, fig. 94. 1876 Patella tasmanica T.-Woods, Proc. Roy. Soc. Tasm., p. 157. 1891 Patella (Scutellastra) ustulata) Reeve, Pilsbry, Man. Conch., vol. 13, p. 101, pl. 22, figs. 11, 12. 1891 Patella (Scutellastra) aculeata Reeve, Pilsbry, Man. Conch., vol. 13, p. 100, pl. 25, figs. 20, 21; pl. 62, figs. 11, 12. 1906 Patella hepatica, Verco, Trans. Roy. Soc. S. Aust., vol. 30, p. 207. Non P. and G. 1924 Patellanax squamifera Reeve, Iredale, Proc. Linn. Soc. N.S. W., vol. 49, p. 239. 1955 Patellanax peroni Blainville, Macpherson, Proc. Roy. Soe. Vict., vol. 67, pt. 2, pp. 232, 233, text figs. (shells and radula). 1957 Patellanax peronii Blainville, Cotton, South. Aust. Moll, Archaeogast., Govt. Print., Adelaide, p. 290, text fig. 194. Records—Southern WESTERN AUSTRALIA; King George Sound (type locality). SOUTH AUSTRALIA; Marino. VIC- TORIA; near Port Phillip Heads. TASMANIA; Port Arthur; Blackman’s Bay. NEW SOUTH WALES; Merimbula; Port Jackson; Cronulla; Shellharbour (all AWBP coll.). Patella chapmani Tenison-Woods, 1875 (PI. 111; Pl. 113, fig. 3) Range—Southern half of Australia and Tas- mania. Remarks—This is a small white limpet that assumes different outlines, varying from irreg- ularly-lobed to a regular 8-pointed star. This latter form, more common in New South Wales is the “Acmaea_ saccharina var. perplexa” of Pilsbry, 1891. It lives in the lower littoral zone among algae on rock platforms and boulders. INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella = 137 Plate 111. Figs. 1-4. Patella (Scutellastra) chapmani Tenison Woods, 1875. Figs. 1-3. Kurnell Botany Bay, New South Wales, 22.5-25 mm., AWBP coll. 19573. Fig. 4. North Har- bour, Port Jackson, New South Wales, 31 mm., AWBP. coll 45675. Description—Shell small, average length 20 mm. (4 of an inch) but grows up to 30 mm. (1% inches), irregularly to regularly star-shaped, with eight prominent radial ribs that project to a vary- ing extent; irregularly and weakly lobate in typ- ical chapmani, regularly and strongly projecting and narrowly-pointed in the form perplexa. Sur- face sculptured with numerous radial, weakly- scabrous lirae, the centre one down each of the eight radial folds being stronger than the rest. Colour of exterior buff, with scattered flecks of light-brown, and often, especially in the perplexa form, with a narrow reddish brown line down the crest of each of the eight radial folds. Interior porcellanous-white, without a clearly defined spatula. Radula—Formula 3 + 1] + (2+1+2) + 1 + 3. Cen- tral teeth five, the middle one only half the size of the other four, lateral massive with four cusps, and the three marginals long and slender, each with a weak blunt cusp. Measurements (mm.)— length width height 30.0 28.0 12.0 North Harbour, Sydney 28.5 27.0 6.0 Kurnell, N.S. W. 23.0 19.5 6.0 Kurnell, N.S. W. 18.5 16.0 4.0 Kurnell, N.S. W. [01-661 | 138. Subgenus Scutellastra A.W. Plate 112. Geographical distribution of Patella (Scutellastra) chapmani Venison Woods. Synonymy— 1873 Patella octoradiata Hutton, Cat. Mar. Moll. N. Z., p. 44 (erroneously attributed to New Zealand). (non Gmelin). 1875 Patella chapmani Tenison-Woods, Proc. Roy. Soc. Tas., p. 157. 1876 Acmaea alba Tenison-Woods, Proc. Roy. Soc. Tas., p. 155. 1891 Acmaea saccharina perplexa Pilsbry, Man. Conch., vol. 13, p. 50, pl. 36, figs. 69, 71. 1915 Patelloida perplexa Pilsbry, Iredale, Trans. N. Z. Inst., vol. 47, p. 430. 1922 Patella perplexa Pilsbry, Peile, Proc. Malac. Soc., Lond., vol. 15, p. 16, text fig. 4. 1924 Patella perplexa Pilsbry, Iredale, Proc. Linn. Soc. N.S. W., vol. 49, p. 238. 1924 Scutellastra chapmani Tenison-Woods, Oliver, N. Z. J. Sci. Tech., vol. 7, p. 244 (radula). 1955 Patellanax chapmani Tenison-Woods, Macpherson, Proc. Roy. Soc. Vict., vol. 67, pt. 2, p. 231, text figs.; pl. 8, figs. 1, 2. 1959 Patellanax alba Tenison-Woods, Cotton, S. Aust. Moll., Archaeogast., Govt. Print., Adelaide, p. 288. Types—The types of chapmani and of alba are in the Tasmanian Museum, Hobart, and that of perplexa in the Academy of Natural Sciences of Philadelphia. Records—AUSTRALIA: NEW SOUTH WALES; Angourie, north coast; North Harbour, Port Jackson; Kurnell, Botany Bay; Shellharbour; TASMANIA (type locality of chapmani); (all AWBP coll.). SOUTH AUSTRALIA: VICTORIA and south- em WESTERN AUSTRALIA (Cotton, 1959). Patella hamiltonensis (Chapman and Gabriel, 1923) Range—Muddy Creek, upper beds, Victoria, Australia, Kalimnan, lower Pliocene. Remarks—This species, described as an acmaeid, and compared with “Patelloida perplexa Pilsbry” by its authors, was recently referred to Patellanax by Darragh, 1970. The present writer has not seen the type material but accepts Darragh’s location B. Powell Patellidae of the species in the Patellidae. The original de- scription follows, but the original accompanying illustration is not clear enough for copying. Description—“Shell solid, irregularly — oval, strongly ribbed; apex sub-central, much eroded and probably originally smooth. The sculpture consisting of about ten rather prominent radiat- ing ribs, the interspaces of which are occupied by finer riblets of varying strength. About three. irregular growth stages are discernible on the shell surface, which are marked by slight over- lapping or sulcation. The area between the major ribs, depressed or fluted, resulting in an undulose margin to the shell. Colour pale ochre.” “Observations—This species approaches Patel- loida perplexa Pilsbry, but differs in the ribs being less pronounced and not salient at the margins.” [Acmaea saccharina var. perplexa Pilsbry, 1891, is actually a synonym of Patella (Scutellastra) chapmani Tenison-Woods, 1875]. Measurements (mm.)— length width height 13.0 12.0 5.0 holotype Plate 113. Fig. 1. Patella (Scutellastra) laticostata Blainville, southern Western Australia. Radula, from Macpherson, 1955, p. 235, text fig. Fig. 2. Patella (Scutellastra) peronii Blain- ville, Victoria. Radula, from Macpherson, 1955, p. 233, text fig. Fig. 3. Patella (Scutellastra) chapmani Venison Woods (as P. perplexa Pilsbry), New South Wales. Radula, from Peile, 1922, Proc. Malac. Soc., vol. 15, p. 16, fig. 4. [01-662] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 139 Synonymy— 1923 Patelloida hamiltonensis Chapman & Gabriel, Proc. Roy. Soc. Victoria, vol. 36 (N. S.), pt. 1, p. 24, pl. 1, fig. 3. 1970 Patellanax hamiltonensis: Darragh, Mem. National Mus. Victoria, vol. 31, p. 173. Types—The holotype and two paratypes are in the National Museum of Victoria, Melbourne, Australia. Patella cooperi (Powell, 1938) (Pl. 114) Range—New Zealand, lower Miocene. Remarks—The_ species belongs to the wide ranging Indo-Pacific flexuosa group, but just how closely related is this Miocene species, cannot be determined on the basis of the only known ex- amples, both of which are in an eroded and badly damaged condition. Description—Shell of moderate size, 50 mm. (2 inches) or more in length, solid, depressed, elon- gated, star-shaped, with eight principal radial ribs that are carinated, very prominent, and strongly projecting at the margin. In the inter- stices there are from 3 to 5 secondary ribs. Apex estimated at about the anterior third. Measurements (mm.)— length width height 50.0 38.0 9.0 (approximately); holotype Synonymy— 1938 Scutellastra cooperi Powell, Trans. Royal Soc. N. Z., vol. 68, p. 379, pl. 39, figs. 13, 14. Records—NEW ZEALAND: Motuihi Island, south coast, Auckland, in conglomerate, basal Waitemata Group, Otaian Stage, lower Miocene. Types—Holotype and paratype in the Auckland Museum. Plate Ll4. Patella (Scutellastra) cooperi (Powell, 1938), New Zealand, Motuihi Island, Auckland, Otaian, lower Miocene. Fig. 1. Holotype, 50 mm., Auck. Mus. 706. Fig. 2. Paratype, 69 mam... Auck. Mus. [01-663] 140) Subgenus Scutellastra A. W. B. Powell Patellidae a a a a ee [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence. | [01-664] November 27, 1973 Subgenus ANCISTROMESUS Dall, 1871 Type: Patella mexicana Broderip and Sowerby, 1829 This is the largest known patellid limpet which sometimes attains a length of 14 inches. It is the only known true patellid living on the west coast of the Americas, with the exception of the sub- antarctic Nacella and Patinigera, of which the latter extends up the Chilean coast as far north as Valparaiso. Dall’s subgenus is based largely upon the dif- ferent form of the branchial lamellae, which are not semicircular as in other patellids, but are produced, twisted and elongated, having an arbor- escent appearance. Also, unique among patellids, the entire animal is black, more or less marbled and streaked with white. The radula is similar to that of other Indo- Pacific Patella except that the median central tooth is fully developed so that there are five evenly-developed centrals in one horizontal series. In other Indo-Pacific Patella species the median central tooth is much smaller, vestigial, or occa- sionally absent, as it is in the European Patella vulgata. Undoubtedly there is some relationship between Ancistromesus and large Indo-Pacific patellids, such as kermadecensis Pilsbry and tucopiana INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 141 Powell. It is assumed that Ancistromesus owes its presence in the tropical West American fauna, as an extreme Indo-Pacific outlier, having reached there in the geological past when the ancient Tethys Sea offered a free equatorial water-way around the globe. Synonymy— 1871 Ancistromesus Dall, American Journal of Conchology, vol. 6, part 3, p. 266. Type, by monotypy, Patella mexicana Broderip and Sowerby, 1829. Patella mexicana Broderip and Sowerby, 1829 (PI. 115) Range—Gulf of California to Peru. Remarks—Apart from its solidity and huge adult size, mexicana has a consistently narrowly ovate outline, and young shells are at once recognised by the presence of a broad, thin, semitransparent margin that extends out abruptly from the outer edge of the porcellanous interior. It is almost certain that Patella gigantea Les- son, 1831, described from a shell “thrown on the coral rocks” at Borabora, Society Islands, is a mexicana that was probably taken there and discarded by a sailor from one of the many whalers that frequented the area at about the time. Apparently Lesson’s shell has never been figured, and enquiries concerning the existence or not of the type specimen were abortive, since at the time of writing, the molluscan collections of the Museum National d’ Histoire Naturelle, Paris, were under general reorganisation. Dr. Harald A. Rehder of the National Museum Plate 115. Patella (Ancistromesus) mexicana Broderip & Sowerby, 1829. Mazatlan, West Mexico, 107-116 mm., AWBP coll. 52692. The world’s largest limpet, known to attain a length of 355 main. — 14 inches. [01-675] 142. Subgenus Ancistromesus of Natural History, Washington, who has just spent six months of intensive collecting at Bora- bora and neighbouring areas, did not find any large limpets resembling gigantea, and he also is of the opinion that gigantea should be considered a synonym of mexicana (personal communica- tion). The dimensions given by Lesson for his gigan- tea, the equivalent of 7 by 5 inches, match exactly the length-width ratio of a series of mexicana. Also Lesson’s description of the interior of his shell—“L’ interi€ur est lissee, blanchatre, avec le fond rougeatre”, applies to occasional examples of mexicana, the reddish brown staining of the spatula area, occurring when the outer surface has been extensively eroded. Description—Shell massive and very large, 200 to 355 mm. (8 to 14 inches) in length, depressed in its younger stages but moderately elevated in the adult, with the apex subcentral to a little nearer to the anterior end. Outline elongate- ovate, noticeably narrowed at the anterior end, the margin thin, wide, flattened and irregularly corrugated in juveniles, but thickened and mi- nutely crenulated in the adult. Sculpture in young shells consisting of eleven low and broad _pri- mary radials and a varying number of secondary radials in between. Adult shells are almost in- variably eroded, encrusted, or riddled by boring bivalves. Colour dull-white externally and_por- cellanous-white within, sometimes diffused with reddish brown over the spatula. In young shells the broad flat thin margin is semitransparent. Measurements (mm.)— length width height 355.0 — _ Keen, 1958, p. 242 185.0 148.0 ca 60.0 Caleta, Acapulco 158.0 114.0 67.0 Caleta, Acapulco 95.0 75.0 16.0 Acapulco Synonymy— 1829 Patella mexicana Broderip and Sowerby, Zool. Journ., vol. 4, p. 369. 1831 Patella gigantea Lesson, Voy. Coquille, Zool., vol. 2, p. 423. 1841 Patella maxima Orbigny, Moll. Amér. Mérid., p. 482. 1855 Patella mexicana Brod. and Sby., Reeve, Conch. Icon- ica, pl. 1, fig. 1. 1871 Ancistromesus mexicanus Brod. and Sby., Dall, Amer. Journ. Conch., vol. 6, pt. 3. 1891 Patella gigantea Lesson, Pilsbry, Man. Conch., vol. 13, p. 156. 1891 Patella mexicana Brod. and Sby., Pilsbry, Man. Conch., vol. 13, p. 108, pl. 31, figs. 59-62. 1958 Patella (Ancistromesus) mexicana Brod. and_ Sby., Keen, Seashells Trop. W. America, p. 242. A. W. B. Powell Patellidae Patella fuenzalidai Herm, 1969 (Pl. 116) Range—Pliocene of northern Caldera Province, Atacama, North Chile. Remarks—This species differs from the Recent mexicana in being more broadly ovate, much finer sculptured, with the primary radials almost obsolete, and in having a much narrower mar- gining bevel. Description—Shell massive and very large, 188-209 mm. (74-84 inches) in length, broadly ovate, only slightly narrowed towards the anterior end, and rather elevated. Sculpture very densely and finely radially lirate, and obsoletely eight- rayed, the rays visible only by the flattened planes between them. Apex a little anterior to the middle. Interior with a large well defined spatula, and a relatively narrow bevelled margin at the perimeter of the shell. Measurements (mm.)— length width height 209.0 187.0 68.0 — holotype 188.0 161.0 61.0 — paratype Synonymy— 1969 Patella (Ancistromesus) fuenzalidai Herm, Zitteliana, vol. 2, p. 131, pl. 14, figs. 1-3. Types—The location of the type is unknown to US. Plate 116. Patella (Ancistromesus) fuenzalidai Herm, 1969. South America, northern Caldera Province, Atacama, North Chile, Pliocene. Holotype, 209 min., and paratype, 1S8 mm., from Herm, 1969, Zitteliana, 2, pl. 14, figs. 1, 2. [01-676] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Helcion 143 Genus Helcion Montfort, 1810 Type: Helcion pectunculus (Gmelin, 1791) This genus has a radula identical with that of Patella, but the gill cordon, unlike that of Patella, is interrupted in front by the head as it is in Cellana. The radula in Cellana differs from that of both the above genera in consisting of a pair of central teeth followed by a pair of laterals. The shell of typical Helcion is cap-shaped, high- arched, with the incurved apex almost at the anterior end. The sculpture consists of radial scaly ribs. Synonymy— 1810 Helcion Montfort, Conchyliologie Systematique et Classification Méthodique des Coquilles, vol. 2, pp. 62, 63. Type, by monotypy: Helcion pectinatus Mont- fort, 1810, which is Patella pectunculus Gmelin, 1791. Plate 117. Figs. 1, 2. Helcion pectunculus (Gmelin, 1791), Port Elizabeth, South Africa, 28 mm... AWBP. coll. 11853. Figs. 3-6. Helcion (Ansates) pellucidus (Limiacus, 1758). Figs. 3, 4. South coast, England, 14 mm., Auck. Mus. 19409. Vigs. 5, 6. (laevis form), 20-24 mim., south coast England, AWBP coll. 52497. Figs. 7, 8. Helcion (Patinastra) pruinosus (Krauss, 1848), Algoa Bay, South Africa, 23-25 mm., AWBP coll. 2910. Figs. 9. 10. Helcion (Patinastra) dunkeri (Krauss, 1848), Port) Alfred, South Africa, 15-20 mim., AWBP. coll. 52495. [01-651] Helcion pectunculus (Gmelin, 1791) (Pl. 117, figs. 1, 2; Pl. 118, fig. 1) Range—South Africa. Remarks—This is a common intertidal species. Description—Shell rather small, up to 25 mm. (over 1 inch) in length, solid, roundly ovate, high arched, with the apex incurved and almost at the anterior end. Sculptured with numerous scaly to spinose ribs, both primary and second- ary. Colour buff to light brownish with the ribs black; interior varying from dull leaden to orange- brown. Often the black radials are interrupted, resulting in a tessellated pattern. Measurements (mm.)— length width height 28.0 24.0 13.0 Port Elizabeth 23.5, 19.5 9.0 Port Elizabeth Synonymy— 1778 Patella pectinata Linn., Born, Index Rerum Natur. Mus. Caes. Vind., p. 441. Plate 11S. Fig. 1. Helcion pectunculus (Gmelin), “Red Sea”, in error ~ South Africa. Radula, from Thiele, 1S91, pl. 28, fig. 23. Pig. 2. Helcion (Patinastra) pruinosus (Krauss), Cape of Good Hope. Radula, from Thiele, 1891, pl. 28, fig. 24. Vig. 3. Helcion (Ansates) pellucidus (Linnaeus), Heligoland. Radula, from Thiele, 1S91, pl. 28, fig. 26. 144. Subgenus Helcion A. W. B. Powell Patellidae 1780 Patella pectinata Born, Test. Mus. Caes. Vind., p. 423, pl. 18, fig. 7. (non Linnaeus, 1758). 1791 Patella pectunculus Gmelin, Syst. Nat. ed. 13, p. 3713; based upon Martini-Chemnitz, Conch. Cab., vol. 1, pl. 7, figs. 56, 57. 1810 Helcion pectinatus Montfort, Conchyl. Systém., vol. 2, p. 62. 1848 Patella pectinata Linné, Krauss, Stidafr. Moll., Stutt- gart, p. 57. 1891 Patella (Helcion) pectinata Linn. Pilsbry, Man. Conch., vol. 13, p. 109, pl. 51, figs. 1-3. 189} Helcion pectunculus Gmelin, Troschel and Thiele, Das Gebiss der Schnecken, vol. 2, pl. 28, fig. 23 (radula). 1948 Helcion pectunculus Gmelin, Stephenson, Ann. Natal. Mus., vol. 11, part 2, p. 278, text fig. 10 (radula). Records—SOUTH AFRICA; Cape of Good Hope (AWBP coll.); Port Elizabeth (AWBP coll.); Port Natal (AWBP coll.); Gouritz River mouth, Cape Province (V. Orr, 1955; Auck. Mus.); Durban (Auck. Mus.). Subgenus ANSATES Sowerby, 1839 Type: Patella pellucida Linnaeus, 1758 This subgenus differs from typical Helcion in being almost smooth. The type species is a sea- weed dweller, which accounts for its compara- tively light build and weak sculpture. Synonymy— 1839 Ansates Sowerby, Conchological Manual, p. 6. Type, by monotypy: Patella pellucida Linnaeus, 1758. 1847 Patina Gray, Synopsis of Contents of British Museum ed. 42, p. 148 (non Rafinesque, 1815). Helcion pellucidus (Linnaeus, 1758) (Pl. 117, figs. 3-6; Pl. 118, fig. 3) Range—Seas of western Europe, from Lofoten Islands, Norway, to Portugal. Remarks—The typical form of this attractive little cap-shaped shell is smooth, and of orange- brown colour, with a few sky-blue longitudinal lines. It lives on Laminaria and Fucus seaweeds, to a depth of about 15 fathoms. The variety laevis is a gerontic form of the species that is thicker, larger, and flattens out towards the margin, where the sculpture is more pronounced. Such shells are always ledged, the early portion being exactly like normal pellucidus. Pilsbry (1891) remarked that the laevis variety is due to station, such individuals being found partly em- bedded in the stems of Fucus. Description—Shell small, up to 24 mm. (1 inch) in length, usually rather thin, ovate, high-arched, the apex immersed and _ situated towards the anterior end. Surface smooth and polished, with very weak radial riblets. Colour golden brown, black tipped at the apex, and usually with a few vivid sky-blue lines running back from the apex to the posterior margin. Measurements (mm.)— length width height 24.0 20.3 10.0 England; laevis form 20.0 15.0 8.0 Pilsbry, 1891, p. 110 14.0 11.0 5.0 S. coast, England Synonymy— 1758 Patella pellucida Linnaeus, Syst. Nat., ed. 10, p. 783. 1777 Patella laevis Pennant, Brit. Zool., ed. 4, vol. 4, p. 125. 1777 Patella intorta Pennant, Brit. Zool., ed. 4, vol. 4, p. 125. 1778 Patella coeruleata da Costa, Brit. Conch., p. 7, pl. 1, figs. 5, 6. 1779 Patella cornea Helbling, Abh. Privatges. Bohm., vol. 4, p. 107. 1803 Patella bimaculata Montagu, Test. Brit., vol. 2, p. 482. 1811 Patella cypridium Perry, Conchology, London, pl. 43, fig. 6. 1813 Patella elongata Fleming, Brewster's Edinb. Encycl., vol. 7 (1), p. 65. 1813 Patella elliptica Fleming, Brewster's Edinb. Encycl., vol. 7 (1), p. 65. 1838 Patella cornea Potiez & Michaud, Gal. Moll. Douai, vol. 1, p. 525. 1891 Helcion (Patina) pellucida L., vol. 13, p. 110, pl. 51, figs. 4-10. 1891 Patina pellucida Linn., Thiele, in Troschel & Thiele, Das Gebiss der Schnecken, vol. 2, pl. 28, fig. 26 (ra- dula). Pilsbry, Man. Conch., Radula—F ormula 3 + 1 + (4) + 1 +3, very similar to the radula of Patella except that the two outer centrals are larger than the inner two; there is no trace of a median central. ? Helcion tella (Bergh, 1871) Range—Sargasso Sea Remarks—This species was described from the soft parts only, of a poorly preserved specimen, the shell of which had been detached and lost. Its author compared his species with the Euro- pean Helcion (Ansates) pellucidus (Linnaeus). Synonymy— 1871 Patina tella Bergh, Verhandl. der k.-k. zool. bot. Ge- sellsch., Wien, 21, p. 1297. 1891 Patina tella Bergh, Pilsbry, Man. Conch., 13, pp. 111- 112. [01-682] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Helcion 145 Subgenus PATINASTRA Thiele in Troschel, 1891 Type: Helcion (Patinastra) pruinosus (Krauss, 1848) Shell very similar to that of Cellana, being low and spreading with the apex varying between the anterior fourth and fifth, but the gill cordon is complete as in Patella. Radula similar to that of Ansates, except for the marginals which are very large. Recent, South Africa. Synonymy— 1891 Patinastra Thiele in Troschel, Das Gebiss der Schnec- ken, vol. 2, p. 325. Type, by monotypy: Patella pruin- osa Krauss, 1848. Helcion pruinosus (Krauss, 1848) (Pl. 117, figs. 7, 8; Pl. 118, fig. 3) Range—South Africa, generally distributed. Remarks—A thin oval shell of low profile, yel- lowish-olive, variously rayed and speckled in dark green or brown, easily distinguished from the next species, dunkeri, by its almost smooth sur- face and radial series of sky-blue spots. Description—Shell rather small, up to 31 mm. (14 inches) in length, but usually about 23 mm. (% of an inch), rather thin, elongate ovate, nar- rowed anteriorly, rather depressed, and with the apex at about the anterior fifth. Sculpture weak, consisting of very numerous narrow — radial thread crossed by dense, much finer, concentric lirae. Colour of exterior yellowish olive sparingly dark-brown speckled and with most of the radials minutely dotted with sky-blue. Interior yellowish olive, shining, without a clearly defined spatula. Measurements (mm.)— length width height 31.0 24.0 9.0 Pilsbry, 1891, p. 113 24.3 19.0 6.0 Algoa Bay 22.0 17.0 5.0 Algoa Bay Synonymy— 1848 Patella pruinosa Krauss, Sudafr. Moll., Stuttgart, p. 56, pl. 3, fig. 9. 1855 Patella pruinosa Krauss, Reeve, Conch. Iconica, pl. 35, figs. 109 a, b. 1891 Patinastra pruinosa’ Krauss, Thiele in Troschel and Thiele, Das Gebiss der Schnecken, vol. 2, p. 325, pl. 28, fig. 24 (radula). 1891 Helcion (Patina) pruinosa Krauss, Pilsbry, Man. Conch., vol. 13, p. 113, pl. 51, fig. 11; pl. 13, figs. 68, 69. 1931 Patella pruinosa fuscoradiata Turton, Mar. Shells Port Alfred, p. 171, pl. 40, fig. 1216. 1932 Patella dunkeri approximata Turton, Mar. Shells Port Alfred, p. 170, sp. 1210. 1948 Helcion pruinosa Krauss, Stephenson, Ann. Nat. Mus., vol. 11, pt. 2, p. 278, text fig. 10 (radula). Records—SOUTH AFRICA: Cape of Good Hope (type locality); Cape Peninsula (AWBP coll.); Port Alfred (Turton, 1932). Helcion dunkeri (Krauss, 1848) (Pl. 117, figs. 9, 10) Range—South Africa, Natal. Remarks—A_ thin oval shell, smaller than pruinosus, and differing from that species in being strongly radially ribbed and_ variously rayed with pink, red, or green but never with sky-blue spots. Description—Shell small, up to 19.5 mm. (% of an inch) in length, very thin, subpellucid, ovate, narrowed anteriorly and_ rather depressed. Sculpture consisting of very numerous fine radi- ating lirae, with linear interspaces, the whole crossed by finer and more dense concentric threads. Colour variable, pinkish-white or pale green, radiately lined in pink, red or greenish- brown. Interior shining with the external pattern showing through; spatula indistinct, yellowish to greenish. Measurements (mm.)— length width height 19.5 13.0 6.75 Port Alfred 17.0 11.0 4.6 Pilsbry, 1891, p. 148 15.0 11.0 4.0 Port Alfred Synonymy— 1848 Patella dunkeri Krauss, Stidafr. Moll., Stuttgart, p. 55, pl. 3, fig. 14. 1855 Patella dunkeri Krauss, Reeve, Conch. Iconica, pl. 38, figs. 124 a, b. 1891 Helcioniscus dunkeri Krauss, Pilsbry, Man. Conch., vol. 13, pl. 16, figs. 11-14. 1932 Patella conspicua Philippi, Turton, Mar. Shells Port Alfred, p. 168, sp. 1196 (in part). 1932 Patella dunkeri formosa Turton, Mar. Shells Port Al- fred, p. 170, pl. 40, fig. 1211. 1932 Patella testudinaria’ Linn., Turton, Mar. Shells Port Alfred, p. 170, sp. 1212. 1932 Patella rufanensis Vurton, Mar. Shells Port Alfred, p. 171, pl. 40, fig. 1213. 1932 Patella) gemmula Turton, Mar. Shells Port Alfred, p. 171, pl. 40, fig. 1214. 1942 Helcion dunkeri Krauss, Tomlin and Stephenson, Proc. Malac. Soc., Lond., vol. 25, pt. 1, pp. 7, 8. 1948 Helcion dunkeri Krauss, Stephenson, Ann. Nat. Mus., vol. 11, pt. 2, p. 278, text fig. 10 (radula). Records—SOUTH AFRICA: NATAL: Wahlberg (Krauss, 1845; type locality); Port Alfred (AWBP coll.). [01-687 | 146 Subgenus Patinastra A. W. B. Powell Patellidae | These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence. | [01-688] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana = 147 Subfamily Nacellinae Thiele, 1929 This subfamily comprises Cellana and Nacella, with its subgenus Patinigera. The chief diag- nostic character is in the radula. In the Patel- linae it is relatively short and folded back upon itself, but in Cellana, and in some species of Nacella (Patinigera), it is very long, sometimes five times the length of the shell, and coiled in a spiral of several loops. In both Nacella and Cell- ana the form and arrangement of the teeth is markedly different from that of Patella. In Patella there are 4 or 5 central teeth, the median one being absent, vestigial or fully developed. The lateral is large with several prominent cusps, and the three marginals are slender, weakly-cusped and probably functionless. On the other hand, in Cellana there is a pair of closely-spaced, long, narrow centrals, alternating with a widely-spaced pair of similar laterals. Between the paired cen- trals there is a vestigial median plate. The three marginals are as in Patella. Both the centrals and the marginals rise vertically almost at right angles to their respective bases. The gill cordon is interrupted by the head in Cellana, but is continuous in Nacella and its sub- genus Patinigera. One feature, the epipodial fringe, is present only in Nacella and Patinigera. The shell in the Nacellinae tends towards semi- translucence, is usually highly glazed to iridescent within in Cellana, but in Nacella (Patinigera), it has a bronzy internal lustre. The genus Cellana is restricted to the Indo- Pacific, except in the New Zealand area, where relict populations extend down into the subant- arctic. On the other hand Nacella and its sub- genus Patinigera’ are exclusively cold water inhabitants of Antarctic and Subantarctic waters. Genus Cellana H. Adams, 1869 Type: Cellana cernica (H. Adams, 1869) Shell of similar shape to that of Patella but the interior is usually highly glazed and _ iridescent. The radula differs markedly from that of Patella but is very similar to that of Nacella. It consists of long recurved pairs of centrals, alternating with similarly-shaped pairs of laterals, usually with an incipient or vestigial median functionless central plate. The three marginals are very weak and functionless also. The whole radula is very much longer than in Patella, sometimes five times the length of the body when straightened out. It lies to the right side of the body where it forms a spiral of up to four double coils. A feature of the animal is the discontinuity of the gill cordon, which is interrupted by the head, unlike both Patella and Nacella in which the gill cordon is complete. The genus is mainly confined to and widely distributed in the Indo-Pacific, ranging from Natal up the east coast of Africa to the Persian Gulf and Arabian Sea, then eastward along the Asiatic coasts to as far north as Japan, the islands of the Indian and Pacific Oceans to the Hawaiian Islands, Society Islands, Juan Fernandez, off the coast of Chile, Australia and New Zealand, in- cluding its subantarctic islands to as far south as Campbell Island. Authentic fossil records for Cellana date back to the lower Miocene of both Australia and New Zealand, and carpentariana from the Northern Territory of Australia, which looks very like a Cellana, could extend the genus back to the lower Cretaceous. Synonymy— 1869 Cellana H. Adams, Proceedings of the Zoological So- ciety, London, p. 273. Type, by monotypy: Nacella (Cellana) cernica H. Adams, 1869. IS71L Helcioniscus Dall, American Journal of Conchology, vol. 6, part 3, p. 277. Type, by original designation: Patella variegata Reeve, 1842, which is Patella ca- pensis Gmelin, 1791. Cellana eucosmia (Pilsbry, 1891) (PI. 119) Range—Red Sea and Gulf of Aqaba. Remarks—This species belongs to the radiata series, but is nearer in shape to karachiensis than it is to typical radiata. From karachiensis it differs in sculpture, being finely radially ribbed, with about every fourth primary a trifle larger, and in Plate 119. Cellana eucosmia Pilsbry, 1891. Ras Banas, Red Sea, 35.5—44.0 mim., AWBP coll. 48217. (01-701 148 = Subgenus Cellana A. W. B. Powell Patellidae its coloration of white flecks and dark maculat- tions in the interstices of a nine-pointed star, the rays of which extend to the margin. This is Reeve’s 1854 version of his variegata of 1842, a very different shell, from unknown local- ity, but here considered to be a synonym of radi- ata capensis. The name variegata, however, is not acceptable as of Reeve at either presentation, since there are two prior usages of that name in Patella, one of Rodding, 1798, and the other of Blainville, 1825. Dall (1870) correctly localised Reeve’s 1854 variegata as coming from the Red Sea area, not Australia, as claimed by Reeve. Then in 1891, Pilsbry provided a new name, eucosmia, for the variegata of Reeve, 1854, and cited the following localities for it—“Suez, Red Sea and Gulf of ‘Akaba,’ Japan and Australia.” However, in 1895, in the Stearns “Catalogue of the Marine Mollusca of Japan,” pp. 112, 113, Pilsbry, without rea- sons, switched his eucosmia to cover a very dif- ferent, common Japanese Cellana, even adding that “The species is not known from any locality outside of Japan.” Pilsbry’s 1891 original proposi- tion must stand for the name of the Red Sea Cellana, since it was clearly introduced as a new name for the 1854 variegata of Reeve, bourn out also by the description, based upon Reeve’s 1854 figures. Plate 120. Figs. 1-5. Cellana radiata (Born, 1778). Fig. 1. Mt. Lavinia, Ceylon, 22 mm., AWBP coll. 45269. Figs. 2, 3. Colombo, Ceylon, 25-29 mm., AWBP coll. 224978. Fig. 4. Gigmoto, Catanduanes Island, Philippines, 23 mm., AWBP. coll. 223090. Fig. 5. Fitzroy Island, Queensland, 25.5 mm., (petalata form), AWBP. coll. 45526. Description—Shell rather large for the radiata group, up to 45 mm. (1% inches) in length, ovate, slightly narrowed anteriorly, and of rather low profile, the apex at a little anterior to the middle; anterior slope straight, posterior slope arched. Sculpture consisting of very numerous narrow radial ribs, weakly but densely scaly where crossed by fine concentric lamellae and growth lines. The radials are fairly even, except that about every fourth one is a trifle larger. Colour of exterior pale yellowish brown, with white flecks and dark-brown maculations in the interstices of a pale-brown, nine-pointed star pattern. Interior yellow, with the dark-brown maculations showing through, except for the spatula, which is dark chestnut-brown, usually more or less clouded with white callus. Measurements (mm.)— length width height 44.0 35.0 12.0 Ras Banas, Red Sea 40.0 31.0 11.0 Ras Banas, Red Sea 35.5 28.5 12.55 Ras Banas, Red Sea 30.0 24.0 9.0 Berbera, Gulf of Aden Synonymy— 1854 Patella variegata Reeve, Conch. Iconica, pl. 16, figs. 36 a-c. “Australia,” in error. (non Reeve, 1842, Conch. Syst., 2, pl. 136, fig. 1). 1870 Patella variegata Reeve (1854), Fischer, J. de Conchyl., 18, p. 167. Suez, Egypt. 1870 Helcioniscus variegatus Reeve, Dall, Amer. Journ. Conch., 6, p. 277, pl. 16, fig. 27 (radula); locality cor- rected to Red Sea and Gulf of Aqaba). 1891 Helcioniscus eucosmia Pilsbry, Man. Conch., vol. 13, p. 148, pl. 71, figs. 61-64 (non Pilsbry, 1595, Cat. Mar. Moll. Japan, p. 112, pl. 7. figs. 7-10; Japan). Records—RED SEA; Gulf of Aqaba (Dall, 1870); Ras Banas (AWBP coll.); Berbera, Gulf of Aden (USNM). Cellana radiata (Born, 1778) Unfortunately the better-known name for this species, Patella rota Gmelin, 1791, must fall as a synonym of Patella radiata Born, 1778. No local- ity was given for Born’s species, but without doubt, the shell he described (1778) and figured (1780) is the common Indian and Ceylon form of the limpet known as rota. The overall distribution of radiata is East Africa from Natal northward to the Arabian Sea, India, Ceylon, and the Asiatic mainland to south- em Japan, the islands of the Indian Ocean, north- em Australia, the Philippines, Palau Islands, Solomons down to New Caledonia, and east- ward across the Pacific to as far as the Mar- quesas. This widely-distributed Indo-Pacific limpet is an exceedingly variable one, but nevertheless (01-702] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 149 several of the more distinctive variants qualify for consideration as geographical subspecies. Relevant synonymy and locality data are listed separately under the respective subspecies. Cellana radiata subspecies radiata (Born, 1778) (Pl. 67, figs. 8, 9; PI. 120) Range—India, Ceylon, West New Guinea and Philippine Islands. Remarks—In this the assumed typical form of radiata the sculpture consists of numerous, nar- row, approximately-equal, flat-topped, radial rib- lets, with linear interspaces. There are no under- lying radial folds, and the shape is regularly and broadly ovate. The coloration is exceedingly vari- able and of no diagnostic significance. Description—Shell of moderate size, up to 37 mm. (1-7/16 inches) in length, of rather light build, roundly ovate, and with a subcentral nucleus. Sculpture consisting of numerous, reg- ular, narrow, flat-topped radial riblets, separated by linear grooves. Coloration variable, the typ- ical form maculated with 9 to 11 bifid radial streaks in purplish brown, upon a_ yellowish ground, the spatula chestnut-brown, except when varyingly clouded with whitish callus. In some examples the radial streaks are broken up into sparse dashes and chevrons. In forma aster Reeve, 1855, from unknown locality, the purplish brown radial maculations are about nine, are very broad, and they alternate with narrow yellowish inter- spaces. In forma luzonica Reeve, 1855, from the Philippines, there is a bold radiate pattern of irregular black streaks upon a transparent yellow ground; in forma scalata Reeve, 1855, also from the Philippines, most of the radial maculations become forked towards the margin, and in forma petalata Reeve, 1854, from North Queensland, Australia, the radial maculations in most ex- amples join up to form a few broad bands. Measurements (mm.)— length width height 37.0 33.0 14.0 Galle, Ceylon 31.0 26.0 9.0 Galle, Ceylon 29.0 25.5 11.5. Colombo, Ceylon 25.5 20.0 8.0 Fitzroy Id., N. Queensland 23.0 19.0 10.0 Colombo, Ceylon 21.0 17.0 7.0 Philippines (f. luzonica) Synonymy— 1778 Patella radiata Born, Index Revum Nat. Mus. Caes. Vind., p. 443: 1780, pl. 18, fig. 10. 1791 Patella rota Gmelin, Linn. Syst. Nat., ed. 13, 1, p. 3720; based upon Martini-Chemnitz, Conch. Cab., vol. 10, p. 330, pl. 168, fig. 1619. East Indies and (in error) West Indies. 1832 Patella reynaudi Deshayes, Bellanger’s Voy. aux Indes- Orient., Zool., p. 411. Ceylon Atlas, pl. 2, figs. 11, 12. 1854 Patella petalata Reeve, Conch. Iconica, pl. 22, figs. 56a, b. Australia. Dec. 1854. 1855 Patella aster Reeve, Conch. Iconica, pl. 30, figs. 80 a, b. Unknown locality. Jan. 1855. 1855 Patella luzonica Reeve, Conch. Iconica, pl. 31, figs. 86a, b. Luzon Island, Philippines. Jan. 1855. 1855 Patella scalata Reeve, Conch. Iconica, pl. 31, figs. 89 a, b. Philippines. Jan. 1855. 1855 Patella nimbus Reeve, Conch. Iconica, pl. 42, figs. 143 a, b. Unknown locality. May 1855. 1891 Helcioniscus reynardi (sic) Desh., Pilsbry, Man. Conch., vol. 13, p. 130, pl. 66, figs. 94, 95. 1911 Acmaea travancorica Preston, Rec. Indian Mus., vol. 6, p. 39. Travancore, India. 1911 Acmaea bombayana E. A. Smith, Proc. Malac. Soc., London, vol. 9, p. 357, text figs. A-C. Bombay, India. 1911 Acmaea bombayana var. ceylanica E. A. Smith, Proc. Malac. Soc., London, vol. 9, p. 358, text fig. D Galle, Ceylon. Records—INDIA: Bombay (USNM 443304); Vurkalay, Tra- vancore (type of Acmaea travancorica). CEYLON: Colombo (AWBP. coll.); Mt. Lavinia (AWBP coll.); % mile S. W. of Dehiwala Village, Colombo (ANSP 224978); Galle (ANSP and AWBP coll.); SABAH; Labuan (Aust. Mus.). WEST NEW GUINEA; Boensaki Island, off Sowek, Soepiori Island, Schou- ten Islands (AWBP coll.). PHILIPPINES: Gigmoto, Catan- duanes Island (ANSP); Luzon Island (type locality of Patella luzonica). QUEENSLAND: Cairns; Fitzroy Island (both AWBP coll.). Cellana radiata subspecies capensis (Gmelin, 1791) (PI. 67, figs. 10, 11; Pl. 121) Range—East coast of South Africa and Natal north to Zanzibar. Remarks—This subspecies has subobsolete to obsolete radial folds, with a superimposed sculp- ture of radial riblets that are dense, linear-spaced and granulose to scabrous. The dark-brown radial markings tend to run together to form a few large squarish maculations, and the spatula usually has a dark bar across it near the top, except in heavily callused adults when an orange smear takes. its place. Gmelin based his species upon “Argenville Conch. pl. 1, fig. 0” and “Kaemmerer Conch. Rudolfi, figs. 1, 2.”°. The Argenville plates are numbered differently in the several editions of that work, so to avoid confusion, Gmelin’s second ref- erence, that of Kaemmerer, 1786, pl. 2, figs. 1, 2, is here selected as the basis of the subspecies, and this action is in accord with the generally ac- cepted concept of capensis and coincides with the admirable illustrations in Krauss, 1848. Description—Shell of moderate size, up to 39 mm. (1% inches) in length, rather lightly built, ex- cept in the fully adult; ovate to elongately-ovate, with the anterior end slightly narrowed, depressed [01-703] 150. Subgenus Cellana Plate 121. Figs. 1-6. Cellana radiata subspecies capensis (Gmelin, 1791). Fig. 1. Chukwani Palace, Zanzibar, 35 mm., AWBP. coll. 214134. Figs. 2, 3. Kizimkazi, Zanzibar, 31-37 min., AWBP coll. 40900. Figs. 4-6. Pondoland, South Africa, 24-33 mim., AWBP. coll. 211763b. to moderately elevated, and with the apex vary- ing from subcentral to the anterior third. Sculp- ture consisting of very numerous, fine, linear- spaced, weakly granulose or scabrous radials. Margin finely denticulated. The nine radial folds, so characteristic of some of the other radiata sub- species, are subobsolete to obsolete in capensis. Colour of interior, yellowish with a silvery lustre, an orange to chestnut-brown spatula, and a few broad radial rays of dark purplish brown. Almost invariably the spatula has a dark-brown blob, running in from the right, at just below the con- striction. Fully adult and senile examples often have the spatula obliterated by a whitish callus, and in these theere is usually a smear of bright- orange at the head end of the spatula. Measurements (mm.)— length width height 39.0 29.0 — Pilsbry, 1891, p. 146 37.0 29.0 12.5 S$. W. Zanzibar 33.0 26.0 14.0 Pondoland 28.0 23.5 8.5 Port Edward, Natal Synonymy— 1791 Patella capensis Gmelin, Syst. Nat. ed. 13, p. 3720, based upon Kaemmerer, Conch. Rudolfi, pl. 2, figs. 1,2. A. W. B. Powell Patellidae 1842 Patella variegata Reeve, Conch. Systematica, vol. 2, p. 15, pl. 136, fig. 1. 1848 Patella capensis Gmelin, Krauss, Stidafr. Moll., Stutt- gart, p. 53, pl. 3, fig. 13. 1891 Helcioniscus capensis Gmelin, Pilsbry, Man. Conch., vol. 13, p. 146, pl. 16, figs. 15-17. 1948 Cellana capensis Gmelin, Stephenson, Ann. Natal Mus., vol. 11 (2), pp. 279, 282; text fig. 11 (radula). Records—SOUTH AFRICA; “Cape of Good Hope” (type locality); Coffee Bay, 1 mile S. of Umtata River, Transkei (V. Orr, 1955; ANSP); Port St. Johns, Pondoland (V. Orr, 1955; ANSP): Port Edward, Natal (V. Orr, 1955; ANSP) near Durban (Mrs. N. Prior); ZANZIBAR: Dembiani, 2 miles N. of Kizimkazi (AWBP coll.); Chukwani Palace (ANSP); Mangap- wani (ANSP). Cellana radiata subspecies enneagona (Reeve, 1854) (Pl. 67, fig. 14; Pl. 121) Range— Madagascar, Andaman Islands, Indo- nesia, Philippines and Bonin Islands, Japan. Remarks—The elongated, narrowly ovate shape, nine prominent radial folds, and irregularly in- dented margin, characterise this form or sub- species. Description—Shell moderately large, up to 53 mm. (2% inches) in length but usuaily smaller, narrowly ovate to irregularly angular, with the apex between the center and the anterior third. Sculpture consisting of 9 conspicuous, rounded, radial folds. The entire surface, folds and inter- spaces alike, is crowded with narrow, crisp radial cords that are rendered slightly scabrous by con- centric growth lines. Margin of shell irregular, being strongly projecting at the terminal points of the radial folds, and concave in the interspaces. Colour pattern of interior variable, consisting of radial dark purplish brown streaks upon a pale creamy ground, or the streaks may either ramify towards the margin or break up into tessellated patterns. Usually there are 9 narrow radial areas free from colour pattern, and these correspond to the external folds. Spatula long and narrow, dark- chestnut, sometimes white-callused to a varying degree. Measurements (mm.)— length width height 53.5 43.0 13.5. Jackson Beach, Bonin Islands 39.0 32.0 10.5 Jolo, Philippines 30.0 21.0 5.5 Catbalogan, Philippines 25.0 20.0 6.0 Andaman Islands [01-704] November 27, 1973 Synonymy— 1854 Patella enneagona Reeve, Conch. Iconica, pl. 18, figs. 44a, b. Unknown locality. Dec. 1854. 1855 Patella articulata| Reeve, Conch. Iconica, pl. 33, figs. 97 a, b. Island of ‘Ticao, Philippines. Mar. 1855. 1891 Helcioniscus enneagona Reeve, Pilsbry, Man. Conch., vol. 13, p. 152, pl. 28, figs. 35, 36. 1891 Helcioniscus articulatus Reeve, Pilsbry, Man. Conch. vol. 13, p. 128, pl. 65, figs. 87, 88. 1959 Cellana enneagona Reeve, Oyama and Takemura, The Moll. Shells, vol. 3, Cellana, pl. 2, figs. 6-8.. Types—Three syntypes of articulata are in the British Museum (Natural History). Records—MADAGASCAR: Pointe Ambarionaombi, S. E. of Nossi Be’ (ANSP). Nosy Antsaibory, N. W. Nossi Be’ (ANSP); Nosy Tanikely, 4 miles S. of Nossi Be’ (ANSP); Pte du Cratere, S. W. Nossi Be’ (ANSP); Nossi Iranja, 32 miles S. W. of Nossi Be’ (ANSP). ANDAMAN ISLANDS: Port Blair (AWBP coll.). INDONESIA: Bali (AWBP coll.). PHILIPPINES: Ticao Island (type of articulata); Cabcaben, S. E. Bataan, Luzon Island, rocky shore (ANSP); Iba, Zambales, Luzon (ANSP); Jolo (USNM); Catbalogan, Samar (USNM). JAPAN: Jackson Beach, Bonin Islands (USNM). INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 151 Cellana radiata subspecies orientalis (Pilsbry, 1891) (Pl. 67, figs. 12, 13; Pls. 123 and 124) Range—Indonesia, New Guinea, North Western Australia, southern Japan, Palau Islands, Solomon Islands, New Caledonia, Loyalty Islands, Fiji, Tonga, Samoa and Marquesas Islands. Remarks—This subspecies or form differs from the typical one in the presence of very distinct radial folds that underlie the normal radial sculp- ture. The colour pattern is variable, ranging from buff with sparse dark-brown chevrons, to broad radial dark stripes, each one occupying an inter- space. It is likely that the latter colour form is Patella tessellata Hombron & Jacquinot, 1541, preoccupied, and later renamed Patella hombroni Dautzenberg & Bouge, 1933. Unfortunately I am unable to verify the identity of hombroni since Plate 122. Figs. 1-10. Cellana radiata subspecies enneagona (Reeve, 1854). Figs. 1-3. Jolo, Philippines, 28-39 mim., USNM 245660. Pigs. 4-6. Jamelo Bay, Luzon, Philippines, 27-36 mm., AWBP coll. 48214. Figs. 7, 8. Pointe Ambarionaombi, Nossi Bé, Madagascar, 30-36 mm., AWBP. coll. 48742. Figs. 9. 10. Jackson Beach, Bonin Islands, Japan, 40-53.5 min., AWBP coll. 621911. [01-705] 152. Subgenus Cellana the Lesson types were not available at the time of writing. Description—Shell of moderate size, up to 41.5 mm. (1% inches) in length, rather solid, roundly ovate, and with a subcentral nucleus. The dense linear-spaced radial riblets are superimposed upon an underlying sculpture of bold, distant, radial folds, 11 or more in number, and these strongly corrugate the margin. The coloration is variable. In the typical form the radiating dark-brown lines or streaks tend to anastomose towards the mar- gin, there forming a series of rectangular blotches. In the eudora form the dark markings are small, sparse, often chevron-shaped, and they show through to the interior, which varies from buff to bright lemon-yellow. This form is widely dis- tributed, ranging from Java to Japan and down through the Solomons to New Caledonia. In the form from the eastern extremity of the radiata A. W. B. Powell Patellidae range, Samoa and the Marquesas Islands, the radial lines of the interior tend to run together and form broad, dark-brown, radial maculations, corresponding to the interspaces of the external radial folds. Measurements (mm.)— length width height 41.5 36.0 11.5 9 Marquesas Islands 34.5 30.0 15.0 holotype of orientalis 32.0 97:5 9.0 Tau Id., Samoa 28.0 25.0 12.0 holotype of eudora 24.0) 20.5 7.0 Russell Ids., Solomons Radula—The radula is typical of Cellana, with a very weak and small medio-central vestigial plate between a pair of long, narrow, fully-developed centrals that curve forward tangentially above, and alternate with a pair of broader laterals, that are wider-spaced. Both the centrals and the lat- erals are indented along one edge to form two or three cusps. Plate 123. Figs. 1-10. Cellana radiata subspecies orientalis (Pilsbry, 1891). Fig. 1. Fiji, 34.5 mm. (lectotype of orien- talis), ANSP. Fig. 2. (paralectotype of orientalis), ANSP. Figs. 3, 4. Lifu, Loyalty Islands, 27 mm. (eudora form), AWBP. coll. 45673. Fig. 5. Niuafou Island, Tongan Group, 27 mm., AWBP coll. 48745. Fig. 6. Tau Island, Manua Group, Samoa, 32 mm., USNM. 513368. Figs. 7-9. Lingatu Point, Banika Island, Russell Islands, Solomon Islands, 15-27 mm., Domin. Mus. Fig. 10. Tonga, 23.5 mm., AWBP coll. 25349. [01-706] November 27, 1973 Synonymy— ?1841 Patella tessellata Hombron & Jacquinot, Ann. Sci. Nat., Zool. & Paleont., ser. 2, vol. 16, p. 190 Mangareva. (non O. F. Miller, 1779). 1891 Helcioniscus rota var. orientalis Pilsbry, Man. Conch., vol. 13, p. 146, pl. 72, figs. 76, 77. Fiji. ?1933 Patella hombroni Dautzenberg & Bouge, J. de Conchyl., vol. 77, p. 416 (nom. nov. pro. P. tessellata H. & J., 1841). 1938 Cellana rota: Adam & Leloup, Mem. Mus. Roy. D’Hist. Nat. Belg., vol. 2, pt. 19, p. 12, pl. 2, fig. 3 (shell), text fig. 3 (radula). Java. 1940 Cellana eudora Iredale, Aust. Zool., vol. 9, pt. 4, p. 433, pl. 33, figs. 13-15, Lifu, Loyalty Islands. 1959 Cellana rota: Oyama & Takemura, The Moll. Shells, vol. 3, Cellana, pl. 2, figs. 3-5. Japan. 1964 Cellana rota: Habe, Shells of Western Pacific in colour, vol. 2, pl. 3, fig. 5. Japan. Types—Lectotype, Pilsbry’s figured specimen of orientalis, here selected, and three syntypes, in the Academy of Natural Sciences of Philadelphia. Holotype of eudora in the Australian Museum, Sydney. Records—JAPAN: Amami Islands (Habe, 1964). GUAM (USNM); PALAU ISLANDS: S. E. end of Eil Malk (ANSP). INDONESIA: Java (Adam & Leloup, 1938). N. W. AUSTRA- LIA: Vansittart Bay (Aust. Mus); SOLOMON ISLANDS: Lingatu Point, Banika Island, Russell Islands, high tide on coral rock (Domin. Mus.); coast near Kopiu, southern Guada- canal, on exposed rock platform (Domin. Mus.); Ysabel (Aust. Mus.). NEW CALEDONIA: N. of Toula (AWBP. coll.); La Roche Percée, Bourail (ANSP). LOYALTY ISLANDS: Lifu (type locality of eudora). TONGA: (AWBP coll.); Niuafoou Island (AWBP coll.). FIJI: (type locality of orientalis). MAR- Plate 124. Cellana radiata subspecies orientalis (Pilsbry). Fig. 1. Banika Island, Russell Islands, Solomons. Radula. Fig. 2. Bali, Indonesia. Radula, from Adam and Leloup, 1938, p. 13, fig. 3 (as rota Gmelin). [01-707] INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana = 153 QUESAS ISLANDS: Atuona Bay, Hivaoa Island (H. A. Pils- bry, Pinchot Exped., 1929; AWBP coll.); Taiohae, Nukuhiva Island (H. A. Pilsbry, Pinchot Exped., 1929; ANSP). SAMOA: Tau Island, Manua Group (USNM). Cellana deformis (K. Martin, 1883) Range—Miocene of Tiji Taon, Java. Remarks—This species is inadequately illus- trated by a side elevation only. It shows marked corrugations, similar to those of Cellana radiata subspecies orientalis (Pilsbry, 1891), but until the type material is examined, an exact evaluation of the species cannot be made. Synonymy— 1883 Patella deformis K. Martin, Samml. Geol. Reichs-mus. Leiden, vol. 1, p. 236, pl. 11, fig. 31. Cellana karachiensis (Winckworth, 1930) (Pl. 126) Range—Gulf of Oman to Pakistan. Remarks—This species seems to be nearest to livescens Reeve, 1855 (formerly cernica H. Adams, 1869) from Mauritius which also has 9-10 broad radiate bands of dark reddish brown, upon a yellowish ground, as well as a moderate development of the 9-folds. P. livescens, however, is more elongately ovate, and flatter, with the apex at about the anterior third. Description—Shell rather large, up to 57 mm. (24% inches) in length, broadly ovate, slightly nar- rowed in front, and moderately elevated, with rounded slopes, and a subcentral nucleus; margin smooth to very weakly crenulated. Sculpture of dense, crisp, radial riblets, rendered granulose by concentric growth lines; riblets varying between 120 and 180, with about 20 of them slightly stronger than the rest, and in some examples there is a subobsolete indication of the “9-fold” state, reminiscent of the enneagona subspecies of radiata. Colour of exterior, pale brownish-buff, with eleven broad radiate bands of deep reddish- brown; internally the spatula is chestnut-brown, often clouded over with pale fawn callus, and sur- rounding the spatula is a zone of yellow, merging with silver towards the margin, the external brown pattern showing through; juveniles have a sparse pattern of radiate reddish-brown dashes on a yellowish ground. Measurements (mm.)— length width height 57.0 46.5 20.0 East Pier, Karachi 42.0 36.7 17.5. East Pier, Karachi 41.5 33.0 12.0 Muscat 35.0 29.0 15.0 holotype = + ’ , 5 5 F 154. Subgenus Cellana A. W. B. Powell Patellidae Zo° 40° 60° 80° 1oo* 7 140° 160° 2 180° 160° : 140° : a ] T | for | R.ORIENTALIS 6)? 5 Mecho I Ks / Foun D von re... 9) ae Wn We FORMOSA: R.ENNEAGONA] ——*Y| 20 . MARIANA NE: A@.r i ea | Yoie | orbs Emwete oI he are€ pees 1s. 3 ive q i “ - “Fenspa ' Mafare Pelmyra MALDIVE :. (eh % 7 LY | €:. Christme: °° z —— z 2 = : Se =r I 7 | | Ae | SEYCHELLES 15. oe “Y Sh ele fe ., Joe, of | R-RADIATA |__| Se, Fis is.| “Penge Pe ‘ie = Ln nazar Se i ee ws 3 : New ay | ih cook a : pea ee i nee ee oe eS eRe ee ee eee Se (i =—cauevaonis Gp +——- saaaitatetaiia (outa ea ia ’ TRAC Is Norfolk | | NEW | ZEALAND Morio Theresa 2 ——— 4_|_ | i 1 20 40° 60 80 100 : 120 140° 160° * 180° * 160° or Plate 125. Geographical distribution of Cellana radiata (Born), its subspecies and related species. Fig. 1. Cellana eucosmia (Pilsbry). Fig. 2. Cellana karachiensis (Winckworth). Fig. 3. Cellana radiata (Born). Fig. 4. Cellana radiata sub- Synonymy— 1930 Patella capensis karachiensis Winckworth, Proc. Malac. Soc., London, vol. 19 (2), p. 80. Not figured. Types—Holotype and paratype in the British Mu- seum (Natural History). Records—PAKISTAN: East Pier, Karachi (type locality); oyster rocks, Karachi (Winckworth coll., British Mus. (Nat. Hist.) ); GULF OF OMAN: Muscat (ex. Winckworth; AWBP coll.). Cellana livescens (Reeve, 1855) (Pl. 127) Range— Mauritius. Remarks—This is a moderately large, rather thin-shelled limpet of low elevation, easily recog- species capensis (Gmelin). Fig. 5. Cellana radiata subspecies enneagona (Reeve). Fig. 6. Cellana radiata subspecies orien- talis (Pilsbry). Fig. 7. Cellana pricei Powell (new species). nised by its striking colour pattern of nine broad purplish brown radial bands, separated by narrow golden-yellow rays; there is a whitish spatula, and the whole of the interior is highly iridescent. It is regrettable that cernica, type of the genus Cel- lana, must fall as a synonym of livescens, de- scribed in error from Mazatlan, west Mexico, but in reality the well-known Mauritian limpet. Description—Shell moderately large, up to 51 mm. (2 inches) in length, rather flat, elongately ovate, with a weakly scalloped margin, and a flattened apex at about the anterior third. Sculp- ture consisting of about 100 or more fine, crisp, slightly scabrous radials of varying sizes, and in addition there are the 9 broadly rounded, radial Plate 126. Figs. 1-3. Cellana karachiensis (Winckworth, 1930). Fig. 1. Karachi, Pakistan, 57 mm., AWBP coll. 46136. Figs. 2,3. Muscat, Gulf of Oman, 41 mm., AWBP coll. 46142. Plate 127. Cellana livescens (Reeve, 1855), Mauritius, 34-40 mim., AWBP coll. 25582. [01-708] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 155 Sees folds, similar to those of the enneagona_ sub- species of radiata; although little raised, these folds are quite distinct in all the material exam- ined. Colour of exterior, olive to pinkish grey, with 9 broad, radiate bands of dark purplish brown; internally the spatula is white to bluish- grey, the remaining area with the external pur- plish brown radials showing through strongly; the narrow interspaces, corresponding to the external folds, are bright golden-yellow; the whole highly iridescent. Measurements (mm.)— length width height 51.0 41.0 12.5 Mauritius 39.0 29.0 10.0 Mauritius holotype 36.0 28.0 10.0 Mauritius Synonymy— 1834 Patella novemradiata Quoy and Gaimard, Voy. “Astro- labe,” Zool., vol. 3, p. 346, Mauritius. (non G. Fischer, 1807). 1855 Patella livescens Reeve, Conch. Iconica, pl. 29, figs. 75 a, b. “Mazatlan, Gulf of California,” in error. June 1855. 1869 Nacella (Cellana) cernica H. Adams, Proc. Zool. Soc., London, p. 273, pl. 19, figs. 7, 7a. Barkly Island, Mauritius. 1891 Helcioniscus novemradiatus Quoy and Gaimard, Pils- bry, Man. Conch., vol. 13, p. 146, pl. 30, figs. 55-58. 1891 Helcioniscus cernica Reeve, Pilsbry, Man. Conch., vol. 13, p. 149, pl. 71, figs. 59, 60. 1891 Helcioniscus livescens Reeve, Pilsbry, Man. Conch., vol. 13, p. 152, pl. 73, figs. 99, 100. Types—The type specimens of livescens should be in the British Museum (Natural History) but I have not been able to locate them. Records—MAURITIUS: Barkly Island (type locality); AWBP coll. Hs is o> Plate 128. Cellana pricei Powell, new species, near Apia, Upolu, Samoa. Holotype, 35.6 min. and paratype, 27 mm. Cellana pricei Powell new species (Pl. 128) Range—Samoa and New Hebrides. Remarks—This species is easily recognised by its dark silvery grey to greenish black colour, re- lieved by short marginal white streaks at the extremities of the primary radials. Apparently the species is restricted in habitat to dark volcanic rock. The relationship is with the radiata series, but the shell is sufficiently distinct, particularly in sculpture, to discount the posssibility of it being merely an ecotype. Description—Shell of moderate size, 35.6 mm. (1% inches) in length, ovate, with irregularly cor- rugated margins, depressed, with the apex vary- ing between subcentral and the anterior third. Sculpture rather coarse‘and irregular, of rounded radial ribs, 14 or 15 of them of primary strength, and these project at the margins, slightly more than do the corrugations between them. The whole surface is crossed by dense crisp subla- mellose concentric lirae. Colour of exterior green- ish black, with an elongated white streak towards the margin upon most of the primary radials; interior dark silvery grey, with the spatula dark olive-brown, clouded in part by a bluish white callus. The marginal white streaks of the exterior show through strongly upon the inner surface. Types—The holotype is in the Auckland Museum (TM..1337): Measurements (mm.)— length width height 35.6 28.0) 8.5 holotype 27.0 21.75 7.0 paratype Records—WESTERN SAMOA: Upolu, half-tide near Apia, on black voleanic rocks (L. Price, 1964); Upolu (Col. R. W. Tate, 1920; Domin. Mus. MF. 83). NEW HEBRIDES: reet near hotel, Tanna (W. Ff. Ponder, 1968). Cellana garconi (Deshayes, 1863) (Pls. 129, 130, 132) Range—Island of Reunion and northern Mada- gascar. Remarks—The writer has seen neither the type nor a photograph of this species. The type was not available at the time of writing, owing to reor- ganisation of the collections of the Muséum Na- tional d’Histoire Naturelle, Paris. It is not certain that the type is located in the collections of that institution. However Deshayes’ excellent illustrations show a shell very similar to a common shell from northern Madagascar, the only marked difference being in the position of the apex, which is shown [01-709] 156 Subgenus Cellana A. W. B. Powell Patellidae near central in Deshayes’ figures but is at about the anterior fourth or fifth in Madagascar shells. Deshayes’ shell is tall-conic, but the Madagascar shells are very depressed. However the position of the apex, which is to a great extent governed by the altitude of the shell, varies considerably within many species of limpets, and is therefore of little taxonomic importance. Upon the assumption that the Madagascar shells represent garconi, then that species ap- pears to be related to the Indo-Pacific testudin- aria and represents a western offshoot of that species, just as vitiensis (=sagitata) is an eastern outlier in the Pacific. Plate 129. Figs. 1, 2. Cellana vitiensis Powell, new name pro Patella sagittata Gould, 1846, non Donovan, 1820; Fiji, 34 mm., AWBP coll. 26939. Figs. 3, 4. Cellana garconi (Deshayes, 1863), Nossi B& Madagascar, 31.5 mm., AWBP coll. 257086. Plate 130. Cellana garconi (Deshayes, 1863), Island of Re- union. Original figures from Deshayes, Moll. de L'Tle Ré- union, pl. 6, figs. 11, 12. The main differences in garconi, compared with testudinaria, are the smaller, more lightly built shell, with dense more definite sculpture, the radials being very numerous, and bearing distinct ovate granules; also the shell tends to be narrow in front, and never has the chevroned or tortoise- shell markings of testudinaria. Description (translation from Pilsbry, 1891, I. c.)—Shell “regularly oval, conoidal, the summit elevated, pointed, very slightly directed forward, situated at the front two-fifths of the length. From the apex radiate a great number of very fine, reg- ular, rather equal riblets, which bear long, obtuse granules. The margins are simple and sharp. The interior lined with very bright nacre of a whitish- brown, the central callus quite large, white, sharply defined by the muscle-scar. The shell is thin, semi-transparent, of a uniform brown- blackish, but if held up toward the light, a few rays of a beautiful red become visible.” Measurements (mm.)— length width height 31.0 95.7 7.0 Nossi Be, Madagascar 25.0 19.5 5.0 Nossi Be, Madagascar 23.0 19.0 9.0 type of garconi Radula (Nossi Bé specimen)—Very similar to that of testudinaria, except for the lateral, which is short, broadly arched and expanded above, and very deeply notched. Synonymy— 1863 Patella garconi Deshayes, Moll. de I'lle Reunion, p. 42, pl. 6, figs. 11, 12. 1891 Helcioniscus garconi: Pilsbry, Man. Conch., vol. 13, p. 150, pl. 66, figs. 100, 101. Records—ISLAND OF REUNION (type locality). MADA- GASCAR; south side of Nossi Iranja, 32 mi. S. W. of Nossi Be’ (ANSP Exped. Sept.-Oct. 1960); Pte. du Cratere, S. W. Nossi_ Be” (ANSP); between Ambatoloaka and Madirokely, S. W. Nossi Be“(ANSP). Cellana testudinaria (Linnaeus, 1758) (Pl. 67, figs. 1, 2; Pl. 131; Pl. 132, fig. 1) Range—Andaman Islands to the Ryukyu Islands and to New Caledonia and North Queensland. Remarks—This is a large, solid, broadly ovate, and rather depressed Cellana, externally rather smooth, of brownish slate colour, radially pat- terned in darker brown, and bluish silvery within. This widely distributed Indo-Pacific species is almost invariably associated with dark volcanic rock, and occurs near and below low tide in ex- posed situations. Pilsbry’s Helcioniscus rota var. discrepans proves to be a synonym of testudin- aria. The type material consists of two undersized, [01-710] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana = 157 , & *hlay Plate 131. Figs. 1-3. Cellana testudinaria (Linnaeus, 1758). Figs. 1, 2. Port Tilig, Lubang, Philippine Islands, 73.5 mm., USNM 245683. Fig. 3. Lectotype of Helcioniscus rota var. badly worn examples from unknown locality. An- other synonym is Dunker’s insignis which Fraun- _feld claimed as coming from the Cape of Good Hope, but almost certainly incorrectly. Fraunfeld’s figure looks like testudinaria, but there is no other record of the species from that area. Plate 132. Fig. 1. Cellana testudinaria (Linnaeus), Java. Radula, from Adam & Leloup, Mem. Mus. Roy. D’Hist. Nat. Belg., vol. 2 (19), p. 12. Fig. 2. Lateral tooth of Cellana vitiensis Powell (new name), Fiji. Fig. 3. Lateral tooth of Cellana garconi (Deshayes), Nossi Bé, Madagascar. discrepans Pilsbry, 1891, 29 mm.; an eroded young. testu- dinari from unknown locality. Two related smaller species, also with a pref- erence for dark volcanic rock, extend the range of the testudinaria type of Cellana, to the west- ward in garconi of the Madagascar-Reunion area, and to the eastward in vitiensis, the latter appar- ently being restricted to the Fiji Islands. Description—Shell solid, reaching a large size, up to 90 mm. (3% inches) in length, but most adults about two thirds that size; broadly ovate, of low rounded profile, with the apex at about the anterior third; margin simple and smooth. Ex- ternal sculpture consisting of weak to obsolete, low, narrow, smooth radial riblets. Colour of exterior dark greenish brown, with a radiate pat- tern in dark brown, within the shell substance, and apparent only when the shell is held to the light; the pattern may consist of radial streaks, joined across in a netted pattern, or it may con- sist of bold chevrons; internally the shell is bluish silvery, with the large elongated spatula greyish white to yellowish brown; the margin of the shell has a continuous border in dark-brown, with ter- minal blotches from the internal radial pattern which also shows through faintly, back almost to the spatula. Measurements (mm.)— (all AWBP coll.). length width height 90.0 77.0 33.0 Bongao, Sulu Archipelago 79.0 67.0 27.0 Melanesia 13:5 64.0 18.0 Lubang, Philippines 65.0 56.0 15.0 Raga, New Hebrides 53.0 43.0 14.5. Bataan, Philippines 30.5 24.5 6.5 near Noumea, New Caledonia Radula—All teeth long and narrow; paired [01-711] 158 Subgenus Cellana A. W. B. Powell Patellidae cAROLI oe 7) & ae =a oe os £ 1% be * PHOENIX Is -z « TOKELAU IS. 4 | ’ “Arto, ie {’ [GARCONT Be ty Coe ae VITIENSIS / —_————— es vis |e — bE sali 22 eZ cope |: NEW ZEALAND a | ; | 20 40° O° 80° 100° zoe 140° TH 160° 160° Plate 133. Geographical distribution of Cellana testudinaria (Linnaeus), Cellana garconi (Deshayes) and Cellana vitiensis Powell (new name). centrals each with a long slender, erect, simple- pointed cusp; paired laterals larger than the cen- trals, each with the cusp moderately curved, ending in a sharp point, and with a weak denticle on each side, at the base of the cusp, and at a point just above middle height of the whole tooth; marginals 3, outer two more or less fused basally, and with a rudimentary cusp on the middle one only. Synonymy— 758 Patella testudinaria Linnaeus, Syst. Nat., ed. 10, p. 783. Locality? 1765 “Lepas ou Patelle,” Argenville, Conch. Traité Gen. Coq. de Mer. ed. 2, pl. 2, fig. P. 1798 Patella patera Réding, Mus. Bolten., vol. 2, p. 8. 1825 Patella rumphii Blainville, Dict. Sci. Nat., 38, p. 95. 1854 Patella testudinaria’ Linne~ Reeve, Conch. Iconica. pl. 4, figs. 6 a, b. 1866 Patella insignis Dunker, Verh. Zool.-bot. Gesell., Wien, p. 941. 1868 Patella insignis Dunker, Fraunfeld, Reise der Novara, Zool. Moll. p. 14, pl. 2, fig. 25. 1891 Helcioniscus testudinaria Linne& Pilsbry, Man. Conch., vol. 13, p. 128, pl. 25, figs. 16-19. 1891 Helcioniscus rota var. discrepans Pilsbry, Man. Conch., vol. 13, p. 146, pl. 72, figs. 78-80. 1906 Helcioniscus mestayerae Suter, Trans. N. Z. Inst., vol. 38, p. 322, pl. 18, figs. 7-9. “Stewart Island, New Zealand,” in error. 1938 Cellana testudinaria Linne& Adam & Leloup, Mém. Mus. Roy. D’Hist. Nat. Belg, vol. 2 (19), p. 10; p. 12, fig. 2 (radula). 1959 Cellana testudinaria Linne, Oyama & Takemura, Moll. Shells, vol. 3, Patella-Cellana, pl. 1, figs. 7-10. Types—The figured holotype and the paratype of discrepans are in the Academy of Natural Sciences of Philadelphia, and the holotype of mestayerae is in the Dominion Museum, Welling- ton, New Zealand. Records—ANDAMAN ISLANDS: Port Blair (AWBP coll.). INDONESIA; Java, Amboina, Bali (Adam and Leloup, 1938). SABAH (BORNEO); Marudu Bay, on surf washed rocks at mid to low tide (USNM). NEW GUINEA; Samberbaba, Japen Island (ANSP. Exped. 1956: ANSP). GULF OF SIAM; Koh Chang (USNM): Koh Kut (USNM). SULU ARCHIPELAGO; Bongao Islands (USNM). PHILIPPINES; west coast, Palaui Island, Luzon (USNM); Marivales, Bataan, Luzon (du Pont- Acad. Exped. (ANSP); Iba, Zambales, Luzon (du Pont-Acad. Exped. 1958 :ANSP); Jamelo Bay, Luzon (USNM); Port Tilig, Lubang (USNM); Gigmoto, Catanduanes Island (du Pont-Acad. Exped. 1958 :ANSP); Cuyo Island, Palawan (ANSP). TAIWAN (FORMOSA); (USNM). RYUKYU_ IS- LANDS (USNM): Nase, Okinawa (AWBP coll.) BANKS IS- LANDS; Vanualava (AWBP coll. 186). SANTA CRUZ IS- LANDS; Vanikoro (A. T. Pycroft, 1932, Auck. Mus.). NEW HEBRIDES; Lamap, Mallicolo Island (ANSP); Steepcliff Bay, Pentecost, Raga (AWBP coll.); Aoba (AWBP. coll.); Gaua (A. T. Pycroft, 1932, Auck. Mus.); Vureas (A. T. Py- croft, 1932, Auck. Mus.). NEW CALEDONIA; Baie des Prunes near Noumea (G. & M. Kline, 1958; ANSP): near Amos, N. E. coast, under smooth basalt boulders, in caverns (L. Price, 1969). AUSTRALIA: Queensland: Cairns (AWBP coll.). Cellana vitiensis Powell new name (Pl. 129, figs. 1, 2; Pl. 132, fig. 2) Range—Fiji Islands. Remarks— Unfortunately the name Patella sag- itata Gould, 1846, is preoccupied by the same combination used by Donovan, 1820, in Rees’ En- cyclopaedia of Conchology. Donovan's “P. sag- ittata’ appears in the encyclopaedia against a rather crude figure at the top left of plate 10 in volume 5, and according to Dr. J. D. Taylor of the British Museum (Natural History) there ap- [01-712] November 27, 1973 pears to be no other reference to the name in the text. Nevertheless Donovan’s name qualifies as validly published, and a new name for Gould’s species becomes necessary since no substitute name for the apparently endemic Fijian species is available. The Fijian species has often been mistaken for the young of testudinaria, but examples of that species of comparable size are more elongate, of greater solidity, lie perfectly flat, the radial sculp- ture is weaker and it does not develop granules. A constant feature of vitiensis is the anterior and posterior concavities of the shell margin, so that when the shell is placed upon a flat surface it can be rocked in a fore and aft motion. The large Cellana testudinaria does not seem to occur in Fiji, or elsewhere east of there. Description—Shell rather small, up to 39 mm. (1% inches) in length but usually much smaller, of light build, broadly ovate, and of rather low profile, the apex at about the anterior fourth, and inclined forward; margin thin and_ sharp, without crenulations. There is a broad but slight concavity in the shell margin both anteriorly and posteriorly, and laterally the margin is slightly broadly convex. Sculpture crisp and delicate, consisting of densely-packed linear-spaced rad- ials, all of which are closely granulose. Colour of exterior dark-brown to bluish black, obscurely rayed with pale blue-green tessellations. When held to the light there is an inner pattern of dark reddish brown, composed mainly of anastomos- ing chevrons. Interior leaden-silvery, with a very narrow blackish margin, and chestnut spatula, often with ill-defined outlines. Radula—Similar to that of garconi but the top of the lateral is longer and less arched. Measurements (mm.)— length width height 37.0 32.00 9.0 Fiji 33.5 30:75 11.0 Viti Levu Bay, Fiji 30.0 26.00 7.5 Viti Levu Bay, Fiji Synonymy— 1846 Patella sagittata Gould, Proc. Boston Soc. Nat. Hist., vol. 2, p. 148 (non Donovan, 1820). 1852 Patella sagittata Gould, U. S. Explor. Exped., vol. 12, p. 337, pl. 29, figs. 449 a-c. 1891 Helcioniscus sagittatus Gould, Pilsbry, Man. Conch., vol. 13, p. 130, pl. 65, figs. 89-92. Types—Holotype and three paratypes in the United States National Museum, Washington; USNM. 5839. Records—FIJI_— (type locality); “Mbega’=Beqa — Island ANSP): Viti Levu Bay, N. E. Viti Levu Island, on smooth dark lava rock in the upper tidal zone (W. O. Cernohorsky coll.). INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 159 Cellana grata (Gould, 1859) (Pl. 67, fig. 7; Pl. 134) Range—Japan and Korea. Remarks—This common Japanese limpet has often appeared in the literature under the name of eucosmia Pilsbry, 1891, which name was first applied to a Red Sea shell, and then later misapplied by Pilsbry, 1895, exclusively to a Japanese Cellana. This latter Cellana is a sy- nonym of stearnsii Pilsbry, 1891, which is a more strongly sculptured form of the false ‘ewcosmia’ of Pilsbry, 1895. However, stearnsii also, must be relegated to the synonymy of grata Gould, 1859, which becomes the valid name for this Japanese shell. Gould’s grata is easily recog- nised by its rather narrowly ovate outline, high profile, and prominent sculpture of numerous scaly to tubercled radial ribs. The exterior of the shell is greyish, with dashes and speckles in light to dark reddish-brown. These intermittent radial maculations show strongly in the interior, which also has a clear cut spatula of reddish chestnut, deepening to dark-brown at the edges. Description—Shell of moderate to fairly large size, 30-56 mm. (1-24 inches) in length, ovate to Plate 134. Cellana grata (Gould, 1859), Japan. Fig. Ll Suwanosejima, Osumi, 56 min., USNM 344009. Figs. 2, 3. Nagoya, kKii, 35 and 35.5 mm., AWBP coll. 20276. Fig. 4 Tosa, 34.5 mm. (stearnsii fori), AWBP. coll. 234298. [01-713] 160 Subgenus Cellana rather narrowly ovate and moderately elevated, with both anterior and posterior slopes, slightly to conspicuously arched. Sculpture consisting of very numerous scaly to imbricately tuberculose radial ribs which are variable in development; the form stearnsii having optimum coarse ribbing. Colour of exterior whitish to dull gray, maculated with reddish brown spots and dashes; interior bluish grey or buff, heavily radially maculated with intermittent dark-brown rays and_ spots, the spatula bright-chestnut in the middle, but deepening to a dark-brown clear-cut outer edge. Measurements (mm.)— length width height 56.0 47.0 27.0 Suwanosejima 50.5 40.5 28.0 Matsushima Id., Korea 38.0 29.0 21.0 holotype of stearnsii, Kii 34.0 25.0 16.0 Tosa, Japan 30.0 24.0 14.0 lectotype of grata; USNM 1965. 30.0 23.0 11.5. Misaki, Japan Synonymy— 1859 Patella grata Gould, Proc. Boston Soc. Nat. Hist., vol. 7, p. 161. 1891 Patella (Helcioniscus) stearnsii Pilsbry, The Nautilus, vol. 4, p. 100. 1891 Patella grata Gould, Pilsbry, Man. Conch., vol. 13, p. 135 (unfigured). 1891 Helcioniscus stearnsii Pilsbry, Manual Conch., vol. 13, p. 132, pl. 48, figs. 16-18. 1895 Helcioniscus stearnsii Pilsbry, Cat. Marine Moll. Japan, p. 112, pl. 7, figs. 4-6. 1895 Helcioniscus eucosmius Pilsbry (non Pilsbry, 1891), Cat. Mar. Moll. Japan, p. 112, pl. 7, figs. 7-10. 1959 Cellana eucosmia Pilsbry, Oyama & Takemura, Moll. Shells, vol. 3, Cellana, pl. 3, figs. 1-3. 1959 Cellana_ stearnsii Pilsbry, Oyama & Takemura, Moll. Shells, vol. 3, Cellana, pl. 3, figs. 4-6. 1961 Patella grata Gould, Johnson, Bull. 239, U.S. Nat. Mus. p. 86, pl. 19, figs. 1, 3 (lectotype). 1962 Cellana lucosmia Pilsbry, Kira, Col. Tlust. Shells of Japan, pl. 5, fig. 10. 1962 Cellana dorsuosa torma_ stearnsii Pilsbry, Habe, Col. Mlust. Shells of Japan, pl. 3, fig. 3. 1967 Cellana grata Gould, Habe & Kosuge, Standard Book Jap. Shells in colour, pl. 3, figs. 4, 5. A. W. B. Powell Patellidae Types—Lectotype of grata, selected by Johnson (1961), in the United States National Museum: USNM 1965. Type series of stearnsii in the Acad- emy of Natural Sciences of Philadelphia. Records—JAPAN: north shores of Niphon (lectotype of grata); Kii Province (types of stearnsii); Nagoya, Kii (AWBP coll.); Tosa (ANSP); Suwanosejima, Osumi (USNM); Kom- inato, Kazusa (AWBP. coll.); Minoshima (AWBP); Misaki (ANSP). KOREA; Matsushima Island (USNM). Cellana mazatlandica (Sowerby, 1839) (PI. 67, figs. 17, 18; PI. 135) Range—Bonin Islands, Northwest Pacific Ocean. Remarks—This very large but comparatively thin-shelled Cellana seems to be restricted to the Bonin Islands. It is easily recognised by its tall conical shape, with straight dorsal slopes, con- spicuous spinose radial ribs, pale yellowish brown exterior, sparsely speckled with black, and rich chestnut-brown spatula, within. Unfortunately the well known name for this striking member of the Japanese fauna, nigris- quamata, has to fall as a synonym of Patella mazatlandica, a misnomer, since the species does not occur in the tropical West American fauna. Also Pilsbry’s Patella (Helcioniscus) boninensis is another synonym, being merely the adult of the species. Description—Shell reaching a large size, up to 90 mm. (3% inches) in length, solid but not massive, of moderate height to tall conical, with almost straight slopes; apex anterior third to sub- central. Sculpture of strong, sharply raised, prom- inently tubercled radial ribs; about 38 primary ribs and a varying number of secondary ribs, making a total of between 48 and 55. Colour of exterior pale yellowish brown, deepening towards the margin, the radials with scattered black spots; Plate 135. Figs. 1-3. Cellana mazatlandica (Sowerby, 1839), Chichi Jima, Bonin Islands, Japan. Fig. 1. 61 mm.; Fig. 2. 75 mm.; Fig. 3. 37 mm., AWBP. coll. 204714 (better known as Doninensis Pilsbry, 1891). [01-714] November 27, 1973 interior silvery-gray to cream, or pinkish white, with the spatula deep chestnut- brown, sometimes partly clouded with white callus; crenulated mar- gin tinged with orange- brown. Measurements (mm.) (all A.W.B. Powell coll.)— length width height 90.0 75.0 37.0 Bonin Islands 78.6 64.4 44.0 — Bonin Islands 61.0 51.0 26.0 Bonin Islands 45.0 39.0 21.5 Bonin Islands Synonymy— 1839 Patella mazatlandica Sowerby, Beechey’s Voy. “Blos- som’, Zool. p. 148, pl. 39, fig. 12. “Mazatlan” in error. 1854 Patella nigrisquamata Reeve, Conch. Icon. pl. 2, figs. 3a, b. “Australia”, in error. 1891 Patella (Helcioniscus) boninensis Pilsbry, The Nautilus, p. 79. Bonin Islands. 1891 Helcioniscus boninensis Pilsbry, Man. Conch. vol. 13, p. 131, pl. 66, figs. 1, 2; pl. 67, fig. 3. 1891 Helcioniscus nigrisquamatus Reeve, — Pilsbry, Conch. vol. 13, p. 125, pl. 19, figs. 35, 36; pl. figs. 13-15. (Concepcion, Chile”, in error. 1895 Helcioniscus nigrisquamatus Reeve, Pilsbry, Cat. Mar. Moll. Japan, p. 112, pl. 7, figs. 1, 2. 1895 Helcioniscus nigrisquamatus boninensis Mar. Moll. Japan, p. 112, pl. 7, fig. 3. 1952 Cellana nigrisquamata Reeve, Kuroda and Habe, Check List Ree. Mar. Moll. Japan, p. 44. 1959 Cellana nigrisquamata Reeve, Oyama and Takemura, The Moll. Shells, vol. 3, Cellana, pl. 3, figs. 9-12. Records—BONIN (ANSP); Chichi Jima (USNM). Man. 48, Pilsbry, Cat. ISLANDS: Ogasawara Island Types—Three syntypes of Patella nigrisquamata Reeve, 1854, are in the British Museum (Natural History). Cellana nigrolineata (Reeve, 1854) (PI. 67, figs. 15, 16; Pl. 137) Range—Japan, common and widespread. Remarks—This very attractive species is easily recognised by the orange-stained spatula and by the intricate external pattern of reddish brown 00° 1z0° INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 161 radial ribs and concentric growth lines on a greenish blue ground. Description—Shell large, up to 78 mm. (3 inches) in length, but usually only about two-thirds that size, ovate, with an almost smooth margin, broadly rounded in profile, depressed to rather tall in fully-grown examples, with the apex varying between the anterior third and subcen- tral. Sculpture consisting of slightly raised, nar- row radial ribs, crossed by weak concentric growth lines; between 50 and 60 radials, including intermediates, in fully adult shells. Colour dis- tinctive; externally greenish blue, with the radial ribs and concentric growth lines picked out in reddish brown, or occasionally in black; intern- ally, bluish silvery, with the external rib pattern showing through; spatula ivory-white but more or stained orange-red, dark- chocolate. Measurements (mm.)— less or sometimes length width height 78.0 66.0 29.5. Chiringashima, Japan 58.0 53.5 16.5 9 Fukura Awaji, Japan 56.5 45.5 20.0 Chiringashima, Japan 45.5 36.5 13.0 syntype, British Museum 42.0 32.0 14.5. Nagasaki, Japan Synonymy— 1854 Patella nigro-lineata’) Reeve, Conch. Iconica, pl. 1S, figs. 43 a, b. “Island of Camiguing, Philippines”, probably in error. 1891 Helcioniscus nigrolineatus Reeve, Pilsbry, Man. Conch., vol. 13, p. 133, pl. 13, figs. 48, 49; pl. 14, figs. 71-74. Helcioniscus nigrolineatus Reeve, Pilsbry, Cat. Mar Moll. Japan. Detroit, p. 113. Cellana nigrolineata Reeve, Kuroda and Habe, Check List. Rec. Moll. Japan, p. 44. 1895 1952 1959 Cellana nigrolineata) Reeve, Oyama and ‘Takemura, The Moll. Shells, vol. 3, Patella-Cellana, pl. 1, figs. 5, 6. 1962 Cellana nigrolineata Reeve, Kira, Shells West. Pacitic in Colour, p. 7, pl. 6, fig. 4. 160° 140° G camous 5 ds Pear/ § Hermes | * Guare CAROLINE 0 Helen Rt ~2 & Plate 136. Geographical distribution of Cellana grata (Gould), Cellana mazatlandica (Sowerby), Cellana exarata (Reeve) and : | ‘MARIANA | Pd is A | re + Soipos Ss PHILIPPINE tee is. Midway oe ee oe ra as ee ee oe ee ee ee ee + —— — —— * Woke | yt ye OF got? -¥ MARSHALL IS, + + Wolfe | } ‘ ” Mafuro 7 Palmyra . Mok | “ |° ee 4, 4 “8 Christmas Cellana talcosa (Gould). (01-715] 162. Subgenus Cellana A. W. B. Powell Patellidae Plate 137. Cellana nigrolineata (Reeve, 1854), Chiringashima, Japan, 57-78 mim., AWBP. coll. 52754. Types—Four syntypes in the British Museum (Natural History), of which one, measuring 47.0 X 37.75 X 11.5 mm., is probably the one figured by Reeve, and this I now nominate as lectotype. Records—"Philippines, Island) of Camiguing’=Camiguan (type locality; probably in error); JAPAN: Hirado, Nagasaki Prefecture (Oyama and Vakemura, 1959); Chiringashima (AWBP. coll); Fukura Awaji (USNM);: Sagami Bay (Bishop Mus.); Manazutu) (Auck. Mus.); Minoshima, Wakayama (AWBP coll). Cellana toreuma (Reeve, 1855) (Pl. 138) Range—Japan to Ryukyu Islands, Mariana Is- lands, Taiwan, Hongkong, China, and Philip- pines. Remarks—There is little doubt that toreuma and amussitata represent extremes in sculptural development of a single species. Ino (1935, p. 31) has shown how. size, rib-strength, outline, and other variations, can be accounted for by position in the tidal zone, nature of the substratum, and degree of exposure to wave stress. The predominant form of the species is long and narrow with subparallel sides, and internally, the spatula is long and narrow also. Certain shells from the Philippines (Plate 71, figs. 7, 8) are broadly ovate, but these are extreme indi- viduals in populations that have narrow shells as well. Pilsbry’s Helcioniscus nigrolineatus var. divergens is still another viariant of toreuma, which is of large size, elongate-ovate, but with rounded rather than subparallel sides, and dense, fine, subgranose radial sculpture (Plate 71, fig. 6). Rugosely sculptured shells (Plate 71, fig. 3) are, according to Ino, found towards low water, where rocks have rough surfaces, con- trasted with rock surfaces from higher levels that have been smoothed by erosion. Description—Shell of moderate size, up to 40 mm. (1% inches) in length, lightly built, very de- pressed, apex between the anterior third and fourth, elongate-ovate, narrow, with flattened sides, and tapered to a narrowly rounded front margin; margins weakly crenulated. Sculpture variable, typically consisting of 30-40 moderately strong, narrow, sharply raised primary radial cords, and from 1-3 secondary cords in each interspace, the whole crossed by fine sharp growth lines, that weakly decussate the radial cords, especially the secondary ones. Colour ex- tremely variable; externally, usually greenish or buff, boldly rayed, mottled and blotched with dark reddish brown; internally bluish silvery, the external pattern showing through strongly; spat- ula ill defined, usually diffused chestnut-brown, but white callused at the anterior end. Measurements (mm.)— length width height 40.0 31.0 9.0 Japan; Pilsbry, 1591, p. 135 39.5 98.5 5.8 Nagasaki 36.25 25.0 55 Waki, Satsuma [01-716] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 163 Plate 138. Figs. 1-8. Cellana toreuma (Reeve, 1855). Fig. 1. Nagasaki, Japan, 39 mm., AWBP coll. 45219. Figs. 2, 3. Waki, Satsuma, Japan, 37 & 35 mim., AWBP coll. 204717, Figs. 4, 5. Tlongkong, 24 & 28 mm., USNM. Fig. 6. Synonymy— 1855 Patella toreuma Reeve, Conch. Iconica, pl. 27, figs. 69 a-c. 1855 Patella amussitata Reeve, Conch. Iconica, pl. 30, figs. 83 a, b. 1855 Patella affinis Reeve, Conch. Iconica, pl. 35, figs. 108 a, b. 21855 Patella toreuma var. tenuilirata Carpenter, Proc. Zool. Soc., Lond. “Monterey,” in error. 1891 Helcioniscus toreuma Reeve, Pilsbry, Man. Conch., vol. 13, p. 135, pl. 13, figs. 50-53. 1891 Helcioniscus nigrolineatus divergens Pilsbry, Man. Conch., vol. 13, p. 134, pl. 73, figs. S1-S4. 1895 Helcioniscus toreuma Reeve, Pilsbry, Cat. Mar. Moll. Japan. Detroit, p. 113. 1925 Patella affinis: (synonym of toreuma), Tomlin and Peile, Proc. Malac. Soc., Lond., vol. 16, p. 195. 1925 Patella amussitata Reeve, (synonym of toreuma), Tom- lin and Peile, Proc. Malac. Soc., Lond., vol. 16, p. 198. 1935 Cellana toreuma Reeve, Ino, Bull. Jap. Soc. Sci. Fisheries, no. 37, pp. 31-36. 1952 Cellana amussitata Reeve, Kuroda and Habe, Check List Rec. Mar. Moll. Japan, p. 44. 1952 Cellana toreuma Reeve, Kuroda and Habe, Check List. Ree. Mar. Moll. Japan, p. 44. 1959 Cellana toreuma Reeve, Oyama and Takemura, The Moll. Shells, vol. 3, Cellana, pl. 2, figs. 9-12. 1959 Cellana amussitata Reeve, Oyama and Takemura, The Moll. Shells, vol. 3, Cellana, pl. 2, figs. 13, 14. 1942 Cellana amussitata| Reeve, Yen, Proc. Malac. Soc., Lond., vol. 24, p. 174, pl. 11, fig. 1. Types—The types of affinis, amussitata and toreuma all of Reeve, 1855, are in the British Museum (Natural History), and that of divergens Anatahan Island, Mariana Islands, 44.5 mm. (-divergens Pilsbry, 1891), AWBP coll. 232358. Figs. 7, 8. Malibon, near Manila, Philippines, 35 & 29 mm., USNM 522110. Pilsbry, 1891 is in the Academy of Natural Sci- ences of Philadelphia. The type locality for toreuma cited by Reeve as “Monterey, Cali- fornia,” is erroneous. Records—CHINA: (ex Cuming, Brit. Mus. (N. H.)). JAPAN: Nagasaki (USNM); Waki, Satsuma; Futami, Hyogo. MARI- ANA ISLANDS: Anatahan Island (all AWBP coll.). HONG- KONG (USNM). TAIWAN: (Oyama and Takemura, 1959, pl. 2. figs. 9-14). PHILIPPINES: Malibon, near Manila (USNM.). 100° r \ ¢ BONIN 1s. ¥ wo dima » Morcas dese an ea Mey IO Eee Cae a acid a | y . MARIANA - & Is P) V7 4 Soipon 2 Guar Eniwetok - a. ‘ Aa ’ res [Pe eee es * + Ponape ; : CAROLINE 2 se 3 0 + Helen RE | : SNe lm Ob Plate 139. Geographical distribution of Cellana toreuma (Reeve). [01-717] 164. Subgenus Cellana A. W. B. Powell Patellidae Cellana exarata (Reeve, 1854) (Pl. 67, figs. 4-6; Pl. 140) Range—Hawaiian Islands. Remarks—This well-known Hawaiian limpet is readily distinguished, when not in an eroded state, by its high-conical shape, with straight dorsal slopes, and black external ribbing, with narrow whitish interspaces. Internally the colora- tion is silvery to bluish leaden, and the clear-cut spatula is leaden to black, except when clouded with white callus. Dr. Alison Kay (1969, pp. 1, 2) advocated separation of exarata into three distinct species, diagnosed as follows:— (1) exarata—Shell black, finely sculptured, the radial ribs not extending beyond the mar- gin; foot of animal dark grey, mantle al- most black; mantle appendages short, ex- tending only 5 mm. beyond the edge of the shell. It belongs to the splash zone, on the islands of Oahu, Molokai, Maui and Hawaii. (2) sandwichensis—Shell black, coarsely sculp- tured, the radial ribs extending beyond the margin; foot of animal yellow, mantle grey; mantle appendages long, some extending 20 mm. beyond the edge of the shell. It be- longs to the low tidal zone, over the same range of localities where exarata is found. (3) melanostoma—Shell cream or white, with brown ribs; foot of animal and the mantle bright green. Outlying locations of the Ha- waiian Chain, Necker, Nihoa, Lehua, Gard- ner Pinnacles and parts of Kauai. After examining the excellent range of exarata material in the Bishop Museum, plus extensive material representative of the locations listed following, the writer is of the opinion that sand- wichensis and melanostoma, so far as present evidence goes, do not qualify for higher status than ecotypes of exarata. The juvenile stage, up to 24 mm. in length, in all three forms is identically tessellated in black and white, after which the radial ribs become continuously black, unless defaced by erosion. Most material from the outlying shoals and pin- nacles of the Hawaiian Chain is subject to ero- sion, which reduces the surface of the adult shell to a uniform cream or white. However, in one lot from French Frigate Shoal, several Plate 140. Figs. 1-5. Cellana exarata (Reeve, 1854), Hawaiian Islands. Figs. 1, 3. Kau Waiahukini, Hawai. 47-53 mm.., AWBP. coll. 198947. Fig. 2. Hilo, Hawaii, 35 mm., AWBP coll. 195871. Figs. 4, 5. Gardner Pinnacles, 53 mi., AWBP coll. 200434. [01-718] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 165 adults have retained the black pigmented ribs right through to the fully adult shell. Regarding differences in the coloration of the animal and in the relative lengths of the mantle appendages recorded by Dr. Kay, the writer has found similar variations in the New Zealand Cellana radians, which has a vertical range ex- tending from low water to the lower edge of the splash zone. In the case of exarata versus sand- wichensis it would be interesting to have obser- vations upon limpet animals from mid-tidal loca- tions. The eroded melanostoma form of exarata is strikingly similar to mazatlandica of the Japanese Bonin Islands, and there is little doubt that there is near relationship. The Bonin species has a tessellated juvenile stage also, but the ribbing in the adult stage is tubercular and not continuously coloured, just sparsely speckled, and the apatula is deep chestnut-brown, not leaden to black, as it is in exarata. Description—Shell reaching a large size, up to 83 mm. (34 inches) in length, but usually be- tween 45 and 60 mm., solid, but not massive, narrowly ovate, with a subcentral nucleus, and conical with straight sides. Sculpture consisting of from 46 to 50 bold, sharply raised, flat-topped radials, that are smooth, except were crossed by weak concentric growth striae; margin shallowly corrugated to deeply indented by square-cut cren- ulations. Colour of exterior consisting of plain black radial ribs, with grey or whitish interstices, the apical area only, tessellated with black and white dashes; internally silvery to bluish-leaden, with the dark ribbing showing through; spatula leaden to solid black, but often partly or com- pletely white callused. When the shell is eroded externally that surface becomes whitish or cream coloured, and the corresponding interior is usu- ally diffused with yellow or orange-brown. Measurements (mm.) (all AWBP coll.) — length width height $3.0 68.5 40.0 Gardner Pinnacles 70.0 58.0 34.0 | Molokai Island 64.0 55.5 37.5. Necker Island 42.0 32.0 12.0. Molokai Island Synonymy— 1839 Patella exarata Nuttall, in Jay, Cat. Shells, 3, p. 38 (nomen nudum). “Oregon, California,” in error. 1854 Patella exarata Reeve, Conch. Iconica, pl. 19, figs. 47 a, b. 1854 Patella undato-lirata| Reeve, Conch. Iconica, pl. 23, figs. 59 a, b. “Sandwich Islands.” 1860 Patella sandwichensis Pease, Proc. Zool. Soc., p. 437. 1870 Helcioniscus exaratus Nuttall, Dall, Amer. Journ. Conch., vol. 6 (3), p. 279, pl. 16, fig. 29 (dentition). 1891 Helcioniscus exaratus Nuttall, Pilsbry, Man. Conch., vol. 13, p. 126, pl. 47, figs. 1-3; 6-12. IS91 Helcioniscus melanostomus Pilsbry, Man. Conch., vol. 13, p. 151, pl. 32, figs. 67-69. 1969 Cellana exarata, sandwichensis and melanostoma: Kay, Hawaiian Shell News, vol. 17, no. 4, pp. 1, 2. Types—Three syntypes in the British Museum (Natural History), of which, one measuring 42.5 x 35.0 x 18.0 mm. is evidently the one figured by Reeve and this I now nominate as lectotype. Records—HAWAHAN ISLANDS: OAHU: Haunama Bay; Manaua_ Islands; Mokolea Rock; Kaena Point; Manana: Moku Manu; Popoia Islet. HAWAII: Hilo; Kona; Kau Waia- Plate 141. Figs. 1, 2. Cellana talcosa (Gould, 1846). Fig. 1. Molokai, Hawaiian Islands, 87 mm. AWBP. coll. 195887. Fig. 2. Hawaiian Islands, 56.5 mm.. AWBP. coll. 22915 (better known under its preoccupied name, argentata Sower- by, 1839). [01-719] 166 Subgenus Cellana A. W. B. Powell Patellidae hukini. KAUAI: Haena. MAUI: West Honolua. MOLOKAI: Kaunakokai (all Bishop Mus.). LA PEROUSE PINNACLE. (Tanager Exped., Bishop Mus.). GARDNER PINNACLES (Tanager Exped., Bishop Mus.). NECKER (Tanager Exped.). NIHOA (Tanager Exped. Bishop Mus.). Cellana talcosa (Gould, 1846) (PI. 67, fig. 3; Pl. 141) Range— Hawaiian Islands. Remarks—This very large species of Hawaiian Cellana, long known as Patella argentata Sow- erby, 1839, must take the name of talcosa Gould, 1846, owing to the prior Patella argentata Bosc, 1801. Apart from large size, talcosa is distinguished by its nearly circular outline, broadly rounded, high-arched_ profile, very numerous, rather reg- ular, radial ribs, and distinctive coloration, the exterior, when not encrusted, being reddish chestnut, and the interior silvery, with the scapula) and _— surrounding white. This species is found on exposed rocky shores and outer reefs near the low tide line. Description—Shell solid, very large, up to 106 mm. (4% inches) in length, broadly ovate, almost circular in outline, and roundly low-conical in profile, with the apex subcentral. Sculpture con- sisting of very numerous, narrowly rounded radial ribs, 58-76 primary, and a few secondary riblets in the interspaces of the lower half of the shell; margins finely crenulated. Colour, exter- nally reddish chestnut, but almost invariably encrusted with algae and limy deposit; internally with a large ivory-white spatula, surrounded by a white callused area, and from there to the mar- gin silvery, with the chestnut ribbing of the ex- terior showing through, especially towards the margin. Measurements (mm.)— muscle impression length width height 106.5 94.0) 45.0 Kona, Hawaii 838.0 $1.5 33.0 Molokai $7.0 51.0 37.5 Molokai Synonymy— 1839 Patella argentata Sowerby in Beechey’s Voy. “Blos- som,” Zool., p. 148, pl. 39, fig. 12 (non Bose, 1801), “Valparaiso, Chile,” in error. 1846 Patella talcosa Gould, Proc. Boston Soc. Nat. Hist.. vol. 2, p. 148. Hawaii, Sandwich Islands. 1852 Patella talcosa Gould, U. S. Explor. Exped., Moll. p. 334, pl. 29, figs. 452 a, b. 1854 Patella cuprea Reeve, Conch. Iconica, pl. 8, figs. 15 a, b., “Swan River,” erroneous. 1891 Helcioniscus argentatus Sowerby, Pilsbry, Man. Conch., vol. 13, p. 127, pl. 18, figs. 29, 30; pl. 65, fig. 93. 1969 Cellana talcosa Gould, Kay, Hawaiian Shell News, vol. 17, no. 4, p. 1. Types—The holotype of talcosa is in the United States National Museum, Washington (USNM. 5824). Records—HAWAIIAN ISLANDS: HAWAII: South Point, Kaukalae; Kona; Kau, Waikapuna; Puako. KAUAI: Koloa: MAUI: Hana; Honolua; Keoneio (all Bishop Mus.). MOLO- KAT: outer reefs (AWBP coll.). NIHOA (Bishop Mus.). There seem to be no Recent records of the species from the island of Oahu, but it occurs there fossil in raised coral reef for- mations. Cellana taitensis (Réding, 1798) (Pl. 75) Range—Tahiti, Society Islands, and Pitcairn Island. Remarks—This rather small-sized Cellana_ is lightly built, of ovate outline, moderately ele- vated, closely and = rather regularly radially ribbed, and of dull greenish colour, maculated with numerous intermittent radial dark-brown lines and blotches. It is possibly closely allied to the Lord Howe Island analogia Iredale. The writer is indebted to Dr. Harald A. Reh- der of the National Museum of Natural History, Smithsonian Institution, Washington, for point- ing out (personal communication) R6ding’s ear- lier name for the well known tahitensis (Pease). Réding’s Patella taitensis was cited as coming from ‘Othaheite’ (= Tahiti), and was based upon Favanne, tab. 1, figs. N, N. Despite the crude- ness of Favanne’s illustrations, they suggest the common Cellana of Tahiti rather than Patella (Scutellastra) flexuosa, the only other patellid limpet known to occur in the Society Group. Description—Shell small, 33.5 mm. (14 inches) in length, but usually smaller, of light build, broadly ovate, moderately elevated, and with the apex at about the anterior third; margin thin, minutely crenulated. Sculpture consisting of very Plate 142. Figs. 1-4. Cellana taitensis (Roding, 1798). Figs. 1, 2. Pitcairn Island, 25-30 mm., AWBP. coll. 26869. Figs. 3, 4. Tautira, Tahiti, 24-25 mm., AWBP. coll. 250703. [01-720] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 167 Plate 143. Geographical distribution of Cellana analogia Ire- dale, Cellana craticulata) (Suter) and Cellana — taitensis (Roding). numerous, moderately strong, narrow, radial rib- lets that are deeply cut by closely-spaced con- centric growth lines, resulting in nodulation, es- pecially towards the margin. Colour: externally, varying from dull greenish to greenish white, with a variable radiate pattern in dark-brown, ranging from intermittent radial lines to bold blotches; internally, iridescent dull dark bluish grey, the terminal points of the external ribbing forming a narrow dark border; spatula dark greyish or greenish brown, sometimes clouded with white callus. Measurements (mm.)— length width height 33.5 27.00 11.50 Tautira, Tahiti 29.0 25.00 14.00 Pitcairn 26.0 21575 8.75 ‘Tautira, Tahiti 18.0 15.00 6.00 Tautira, Tahiti n aan | as 6 * PHOENIX 18 ; ". TOKELAU IS. ]: MARQUESAS IS. SAMO4 jo TUAMor Fo GA IS. a U ARCH Soe ern pia *. Te @ u . I A 7 TAITENSIS 4 New rem , TONGA SOK 8 8 is LTAITENSTS | toe ae. 75. CALEDONIA. a NS ie ee, ne ee eee TRAC 1S S&S aa ‘ HE CRATICULATA XK 3 . Norfolk - * ® of 7 [anatocta Oo cacicsce @: Lord Howe ° Synonymy— 1798 Patella taitensis Réding, Mus. Bolten., pt. 2, p. 7, sp. 68. Based upon Favanne, tab. 1, figs. N. N. 1868 Tectura tahitensis Pease, Amer. Journ. Conch., vol. 4 (3), p. 98, pl. 11, fig. 21. 1891 Helcioniscus tahitensis Pease, Pilsbry, Man. Conch., vol. 13, p. 129, pl. 67, figs. 4-8. 1907 Patella (Helcioniscus) tahitensis Pease, Couturier, J. de Conchyl., vol. 55 (2), p. 173. (Pitcairn Island). 1966 Cellana tahitensis Pease, Rehder, Hawaiian Shell News, vol. 14 (8), p. 5. Pitcairn Island. Records—TAHITI (type locality): S. W. of Tautira, on basalt rocks in the splash zone (R. Robertson, 22 July, 1952; ANSP.). PITCAIRN ISLAND (AWBP. coll.): Bounty Bay (Rehder, 1966). TUAMOTU ARCHIPELAGO: Mangareva (Aust. Mus.). Cellana ardosiaea (Hombron and Jacquinot, 1841) (PI. 73, figs. 7, 5; Pl. 144) Range—Island of Juan Fernandez, off the coast of Chile. Plate 144. Cellana ardosiaea (Hombron & Jacquinot, 1541), Island of Juan Fernandez, off Chile, 46-55 mm., AWBP coll. 48228. [01-721] 168 = Subgenus Cellana Remarks—This is the furthest east yet re- corded for the Indo-Pacific genus Cellana. Pils- bry (1891) considered ardosiaea to be allied to the Society Islands taitensis, but that is a most unlikely relationship. From all other species of Cellana, the Juan Fernandez shell stands apart, with its nearly circular, spreading form with its straight dorsal slopes, high conical profile, and its small, erect, nearly central apex. Unfortunately the writer has no preserved animals of ardosiaea, but reference to Schuster (1913) and to Thiem (1917), respectively, leave no doubt that the species is a Cellana, not a Nacella_ (Patinigera), which latter relationship one would have expected, owing to the geo- graphical proximity of ardosiaea to the South American mainland. The epipodial fringe, so characteristic of Nacella and its subgenus Patinigera, is absent in ardosiaea, as also is any trace of the equally characteristic bronzy coloration of the shell. Thiem (1917, p. 389) described in ardosiaea a presumed sensory organ, the “vorderer sub- palliater Sinnestreif” (anterior subpallial sensory stripe), and a longer posterior one, the former evidently the same structure as Fretter and Graham's (1962, p. 118) “lateral glandular streak” in Patella. The anterior sensory stripe, or lateral glandular streak, was noted in several species of Cellana, but not the ‘posterior stripe,’ Plate 145. Figs. 1-5. Cellana conciliata Iredale, 1940. Figs. 1, 2. Lammermore Heads, Keppel Bay, Queensland, 35-38 mm., AWBP. coll. 45417. Figs. 3-5. Keppel Bay, Queensland, 34-39 mm. (Fig. 5, marked X, compared with holotype), AWBP coll. 45423. A. W. B. Powell Patellidae which possibly, could have resulted from con- traction during preservation. Description—Shell moderately large, up to 58 mm. (2% inches) in length, rather solid, broadly ovate, with the small erect apex nearly central; tall-conical with the sides descending almost perfectly straight; margin smooth to weakly cren- ulated. Sculpture consisting of weak, evenly- spaced, radial primary cords, with 3-4 secondary cords or threads in each interspace, the whole crossed by numerous concentric growth threads. Colour of exterior light bluish olive, darker to- wards the margin; the apex yellowish to reddish brown; interior silvery bluish grey, except for the spatula, which is yellowish to orange-brown, and there is a narrow rim of greenish olive at the margin. Radula—The radula, as figured by Schuster (1913, p. 304, text fig. V), is not diagnostic, since there is no basic difference between the radula of Cellana and that of the Nacella group. Measurements (mm.)— length width height 57.5 51.5 23.0 all Juan 57.5 52.0 21.5 Fernandez 55.0 46.0 20.5 Island 46.0 39.0 17.0 37.0 32.5 14.5 Synonymy— 1841 Patella ardosiaea Hombron & Jacquinot, Ann. Soc. Nat., vol. 2 (16), p. 190. 1854 Patella clathratula Reeve, Conch. Iconica, pl. 14, figs. 30 a, b. 1891 Helcioniscus ardosiaeus H. and J., Pilsbry, Man. Conch., vol. 13, p. 124, pl. 32, figs. 63-66. 1913 Helcioniscus ardosiaeus H. and J., Jahrb., Jena, Suppl. 13, pp. 281-384. 1917 Helcioniscus ardosiaeus H. and J., Naturw., vol. 54, pp. 333-404. Records—JUAN FERNANDEZ: (Stearns coll., USNM ANSP). Schuster, Zool. Thiem, Zool. Cellana conciliata Iredale, 1940 (Pl. 145; Pl. 148, fig. 3) Range—North Queensland down to Bargara, South Queensland. Remarks—This species is easily distinguished from tramoserica by its very fine and dense ra- dial sculpture, and in the adult stage by a dis- proportionate broadening of the posterior end. Also the radula differs from that of tramoserica in that all the teeth are shorter and_ rather stouter. Description—Shell of moderate size, up to 40 mm. (1% inches) in length, broadly ovate, with very weakly scalloped edges, and rather low in [01-722] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 height, with the apex subcentral. Sculpture deli- cate and rather even, consisting of very numer- ous radial riblets, due to little difference in strength between primaries and_ secondaries; about 140 riblets in all. As the shell reaches mature size, it tends to gain little height but spreads posteriorly. Concentric growth lines are weak and apparent only at the margin in adults. Colour greenish or bluish grey externally, often broadly or narrowly radially banded in dark brown, with pale chestnut lines in the interstices; young shells often uniformly dark greenish grey; silvery-blue to creamy-white within, sparsely and faintly rayed with bluish grey towards the mar- gin; spatula fawn to dark yellowish brown, often clouded with a white callus in mature shells. Radula—Similar to that of tramoserica but all the teeth are shorter and stouter than in that species (Macpherson, 1955, p. 239). Measurements (mm.)— length width height 44.0 39.0 15.0 Keppel Bay; paratype 40.0 34.0 14.0 Keppel Bay; holotype 38.7 34.0 15.0 Keppel Bay; topotype 33.0 28.0 14.5. Keppel Bay; topotype Synonymy— 1940 Cellana conciliata Iredale, Aust. Zool. 9, p. 432, pl. 33, figs. 1-3, 19, 20. 1955 Cellana conciliata: Macpherson, Proc. Roy. Soc. Vict., vol. 67 (2), p. 238, pl. 10, figs. 1, 2. Types—Holotype and paratypes in the Austral- ian Museum, Sydney; paratypes and topotypes in Powell collection, Auckland. Records—QUEENSLAND: Keppel Bay (type locality); Bargara, near Bundaberg (Mrs. J. Kerslake; AWBP coll.). Cellana turbator Iredale, 1940 (Pl. 146, figs. 5-7; Pl. 148, fig. 4) Range—Caloundra, south Queensland. Remarks—This is a small conical species that differs from the young forms of tramoserica in sculpture, the radial ribbing being coarsely nod- ulose, in coloration, and also in the radula, as described below. Description—Shell small, up to 18 mm. (% inch) in length, regularly ovate, with an elevated, roundly-conical, profile; apex at the anterior third. Sculpture bold, consisting of about 25 nodulose, primary, radial ribs, with a single sec- ondary radial, almost as strong, in each inter- space; margin weakly crenulated. Colour, green- Cellana 169 ish white externally, and creamy, pinkish, or silvery-white within; sparsely and intermittently lined and speckled with dark-brown; spatula dark chestnut, with clear outlines, or diffused with callus. Radula—F ormula 3+1+(1+0+1)+1+3. The radula is distinctive; the two central teeth are sharply pointed but have a small spur on the outer edge; the bicuspid laterals have a lon inner cusp, with a prominent notch about one fourth of the way down from the tip, and a blunt conical basal cusp, rising to a third the height of the main cusp; marginals three, long and_ slender, the outer one the largest (Macpherson, 1955, p. 238). Measurements (mm.)— length width height 20.0 15:5 7.0 Caloundra 18.0 14.7 5.8 Caloundra 15.0 12.0 6.0 holotype 14.5 12.0 5.0 Caloundra Plate 146. Figs. 1-4. Cellana tramoserica (Holten, 1802) Figs. 1, 2. Manly, New South Wales, 34 mm., AWBP. coll. 3995. Figs. 5-7. Cellana turbator Iredale, 1940, Caloundra, Queensland, 14.5—20 mm. (One marked X, compared with holotype), AWBP. coll. 45440 [01-723] 170 Synonymy— 1940 Cellana turbator Iredale, Aust. Zool., vol. 9, p. pl. 32, figs. 16-18. 1955 Cellana turbator: Macpherson, Proc. Roy. Soc. Vic- toria, vol. 67 (2), p. 239, pl. 10, figs. 3, 4. 433, Types—Holotype and paratypes in the Austral- ian Museum, Sydney. Records—Known only from the type locality, Caloundra, Queensland. Cellana tramoserica (Holten, 1802) (Pl. 73, figs. 1-3; Pls. 146-148) Range—South Australia, Victoria, east coast of Tasmania, New South Wales and southern Queensland. Remarks—This species is distinguished from solida by the more numerous, subcarinate, linear- spaced ribs, and resultant finer marginal crenu- lations, the variegated colour pattern, smaller adult size, and lesser solidity. The various colour forms are described below. Iredale’s sontica from Caloundra, South Queensland, is a small rather drab-coloured form of the species, that does not merit separation. Dr. Hope Macpherson (1955, p. 238) remarked that a series covering both typical tramoserica and sontica showed that there were no radular differences between the two forms. Description—Shell moderately large, up to 60 mm. (2% inches) in length, broadly ovate, roundly conical with the apex subcentral, and with a finely scalloped margin. Sculpture consisting of about 36 strong, subcarinate radial ribs with narrow interstices, the whole crossed by dense, fine, sharp growth lines. Colour exceedingly variable: externally yellowish, pink, or light- brown, with some of the primary ribs dark-fawn, or chocolate, either plain or with elongated whitish patches; again, some of the ribs may be reddish, and in others the rib interstices only may be lined in dark-brown; internally the shell is often yellowish to orange, or golden nacreous, with the spatula varying from dark-fawn to a whitish callus; the margin is variously radially lined by the dark, external ribbing showing through the thinner outer edge. Radula—Formula 3 + 1 + (1+0+1) + 1 + 3. The pair of central teeth are long, curved and _ uni- cuspid, as in solida, but the pair of bicuspid laterals have a distinct notch, half way down from the tip, and there is a small conical cusp at Subgenus Cellana A.W. B. Powell Patellidae Alan Plate 147. Cellana tramoserica (Holten), Bay, Great Australian Bight, South Australia. Radula. the base; of the three marginal teeth, the outer two are sharply pointed, but the inner one has the tip bent over slightly to form a blunt cutting point (Macpherson, 1955, p. 239). It is of interest that a radula from a specimen taken at Alan Bay, Great Australian Bight was four times the length of the shell. Measurements (mm.)— length width height 59.5 48.0 28.0 Noosa Head, Queensland 54.0 46.0 24.0 Manly, New South Wales 37.0 31.0 16.0 Long Reef, New South Wales 22.5 19.0 6.0 The Spit, Port Jackson, N.S.W. Synonymy— 1802 Patella tramoserica Holten, Enum. Syst. Conch., Chem- nitz, p. 85 (based upon Chemnitz, Conch. Cab., vol. 11, pl. 197, figs. 1912, 1913.) 1825 Patella variegata Blainville, p. 100. 1825 Patella tramoserica Martyn, Sowerby, Cat. Tankerville Coll., p. 30. Dict. Sci. Nat., vol. 38, 1831 Patella jacksoniensis Lesson, Voy. “Coquille” Zool., vol. 2 (1), p. 418. 1854 Patella tramoserica Martyn, Reeve, Conch. Iconica, pl. 13, figs. 27 a-c. 1891 Helcioniscus tramoserica Martyn, Pilsbry, Man. vol. 13, p. 142, pl. 70, figs. 49, 52. 1924 Cellana variegata ariel Iredale, Proc. Linn. Soc. N.S.W., vol. 49, p. 242. 1940 Cellana sontica Iredale, pl. 33, figs. 10-12. Conch., Aust. Zool., vol. 9, p. 433, [01-724] November 27, 1973 1955 Cellana tramoserica Holten, Macpherson, Proc. Roy. Soc. Victoria, vol. 67 (2), p. 237. Records—SOUTH AUSTRALIA: Alan Bay, Great Australian Bight (I. G. Marrow); Aldinga (AWBP. coll.). VICTORIA, St. Kilda; Mornington; Point Nepean (Macpherson, 1955); Port Fairy (type of ariel). TASMANIA: east coast, rare and small (W. L. May, 1923). NEW SOUTH WALES: Botany Bay (type locality); Long Reef; Manly; The Spit, Port Jackson; Shellharbour. QUEENSLAND: Port Douglas; Point Vernon (Mrs. J. Kerslake; AWBP coll); Noosa Head; Stradbroke Island; Caloundra (type of sontica); near Brisbane (all AWBP coll.). Cellana solida (Blainville, 1825) (Pl. 73, figs. 4-6; Pl. 148, fig. 1; Pl. 150) Range—Tasmania, Bass Strait Islands, and Victoria to eastern South Australia, in the algal zone of the lower littoral. Remarks—This is a large solid limpet, sculp- tured with bold, rounded, radial ribs. Blainville’s solida applies to the smaller size range of the species, and his rubraurantiaca to the fully adult in which the internal colour usually deepens to orange-red at the margin. A conspicuous feature 2 AY) dy a Plate 148. Radulae of Australian Cellana in’ protile; lateral tooth (left) and central tooth (right). Fig. Ll. Cellana solida (Blainville). Fig. 2. Cellana tramoserica (Holten). Fig. 3. Cellana conciliata Iredale. Fig. 4. Cellana turbator Iredale. All from) Macpherson, 1955, Proce. Roy. Soc. Victoria, vol. 67, pp. 236, 238, 239 and 240. INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 171 of this species is the clearly defined dark-coloured spatula, which varies from olive-brown to green- ish or bluish slate. Description—Shell large, up to 79 mm. (over 3 inches) in length, solid, broadly ovate, with a deeply scalloped margin, tall conical, with the apex varying between subcentral and the anterior third. There are about 26 strong, rounded, radial ribs, crossed by dense, sharp-edged growth lines. Colour variable with age; young shells are grey or greenish grey within, the spatula clearly de- fined, olive to bluish slate; becoming yellowish, and finally pale orange at the margin, where bold radiate dark red-brown radials show through the shell substance; exterior dull grey to pinkish buff, often with radiate yellowish brown streaks in the rib interstices. Fully grown examples tend to have the spatula more or less masked by a thick whitish callus, and the margin is bright reddish orange, regularly banded by the reddish brown radials. In this latter form, rubraurantiaca, the exterior is pinkish white. Radula—Formula 3 + 1 + (1+0+1) + 1 + 3. The two centrals are long, slender, curved and _ uni- cuspid; the pair of laterals are bicuspid, the main member similar to the centrals, but with a small additional cusp at the base; the three marginals are narrow and slender, the inner one curved over at the top into a cutting edge, the other two simple (Macpherson, 1955, p. 236). Measurements (mm.)— length width height 78.5 67.0 38.0 Bass Strait 75.0 62.0) 31.0 St. Helens, Tasmania 74.0 61.5 33.0 South Australia 51.0 43.5 19.5. St. Francis Id., S. Australia Ss, fe . ote Fistis oF Lar = Trae | : CALEDONIA ___ “@_ sd | Norfolk | ) | ©] HowENSTS NEW ZEALAND pase 1 Hee 160° 180° ' Plate 149. Geographical distribution of Cellana tramoserica (Holten), Cellana turbator Iredale and Cellana howensis Iredale. [01-725] 172 Subgenus Cellana A. W. B. Powell Patellidae Plate 150. Cellana solida (Blainville, 1825), Corny Point, South Australia, 51 and 42 mm., AWBP coll. 187. Synonymy— 1825 Patella solida Blainville, Dict. Sci. Nat., vol. 38, p. 110. 1825 Patella rubraurantiaca Blainville, Dict. Sci. Nat., vol. 38, p. 111. 1849 Patella limbata Philippi (non Réding, 1798), Abbild. und Beschr. Conch., vol. 3 (6), p. 71. 1854 Patella limbata Philippi, Reeve, Conch. Iconica, pl. 13, figs. 29 a, b. 1891 Helcioniscus limbata Philippi, Pilsbry, Man. Conch., vol. 13, p. 143, pl. 71, figs. 53-56; pl. 17, figs. 28, 29. 1955 Cellana solida Blainville, Macpherson, Proc. Roy. Soc. Victoria, vol. 67 (2), p. 236. 1962 Cellana solida Blainville, Macpherson and Gabriel, Mar. Moll. Victoria, p. 45. Records—VASMANIA: Port Arthur (AWBP coll.); Circular Head (AWBP coll.); Penguin (A. F. B. Hull; AWBP coll.); William’s Island, Bass Strait. VICTORIA: Cape Otway; Wilson’s Promontory (Macpherson, 1962, p. 47). SOUTH AUSTRALIA: Point Sinclair (AWBP coll.); St. Francis Island (AWBP coll.); Corny Point (AWBP coll). ?Cellana carpentariana Skwarko, 1966 (Pl. 152, fig. 1) Range—Australia, Mount Young, Northern Territory, late Neocomian, lower Cretaceous. Remarks—This species bears some resemblance to the Recent enneagona Reeve, 1854 and the Australian lower Miocene cudmorei Chapman and Gabriel, 1923. If carpentariana is really a Cellana then it is the earliest known member of the genus. Description—(original) “The shell is moderately large and inflated. Its apex is obtusely pointed, situated anteriorly, and not incurved. The slopes are straight in the front and on the sides of the shell but convex on the posterior wall, with a wavy posterior margin. The posterior slope is ornamented with four primary, three secondary, and six tertiary straight radial ribs which gradually increase in breadth away from _ the umbo. The primary ribs are straight, sharp- crested, and prominent. Radial ribbing is also : et al ors ie, A . s : i My ig wnew se me f / | EBRIDES ..- | ov. : ; FA. Ce a e\ «| ve Fans. 4 Hees re who oN eee ee we ¢ Se OS oh | CONCILIATA |") eS ; _ Nor o/h ? | \ NEW SOLIDA ZEAE 120° 160° 180° Plate 151. Geographical distribution of Cellana conciliata Iredale and Cellana solida (Blainville). present on the sides, and on the anterior end of the shell, but is not distinct and the costae there seem to belong to one order only. Ribs are crossed by irregular growth-rugae and somewhat more irregular growth striae”. Measurements—No_ size indicated other than “moderately large.” Synonymy— 1966 Cellana (?) carpentariana Skwarko, Comm. Aust. Dept. Nat. Dev. Bur. Min. Res. Geol. and Geophys. Bull. 73, p. 120, pl. 14, fig. 11. Cellana cudmorei Chapman and Gabriel, 1923 (Pl. 152, fig. 2) Range—Australia, polyzoal rock of Batesford, near Geelong, Victoria, Batesfordian Stage, lower Miocene. Plate 152. Fig. 1. ?Cellana carpentariana Skwarko, 1966, late) Neocomian, lower Cretaceous, Mt. Young, Northern Territory, Australia. Holotype, from Skwarko, 1966, pl. 14, fig. Ll. Fig. 2. Cellana cudmorei Chapman & Gabriel, 1923, Batesford, Victoria, Australia, lower Miocene; 40 mm. Holo- type, from Chapman & Gabriel, 1923, pl. 1, fig. 1. (01-726] , November 27, 1972 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 173 Remarks—Chapman and Gabriel considered this species to be ancestral to tramoserica, but it is not related to that species, being in fact a member of the radiata series, as shown by the nine broad primary rays, a very similar Recent shell being radiata subspecies enneagona Reeve, 1854, with its synonym articulata Reeve, 1855, the latter from the Philippines. Similar shells range northward to the Bonin Islands, Japan. This tendency to develop nine primary rays oc- curs sporadically throughout the radiata eries. Description—(original) “Shell large, elevated, oval, rather strongly built; apex situated about one-third from the anterior margin. Sculpture consisting of numerous strong riblets, with two or three smaller one occupying the interspaces. Growth-lines undulate, fine, not well developed.” Measurements (mm.)— length width height 40.0 30.0 10.0 holotype Synonymy— 1923 Cellana cudmorei Chapman and Gabriel, Proc. Roy. Soc. Vict., new ser., vol. 36, p. 23, pl. 1, fig. 1. Types—The holotype is in the National Museum of Victoria. Cellana hentyi Chapman and Gabriel, 1923 Range—Australia, shell bed at Forsyth’s, Grange Burn, near Hamilton, Victoria, Kalimnan Stage, lower Pliocene. Remarks—Chapman and Gabriel compared their species firstly with Patella peronii (as squamifera) and then as an alternative with Cellana’ tramoserica (as variegata). However the latter interpretation is the more likely one, the sculpture being similar to that in the Recent tramoserica, except that the concentric lines produce knotted nodes where they cross the radials. The original illustration is too indistinct to copy. Description—(original) “Shell of medium size, elevated, narrowly oval, apex a little in front of centre. In the present state of fossilisation the apex is denuded of ornament. Surface ornament consisting of moderately strong radiating ribs, with several intermediate, less pronounced rib- lets; these are crossed by growthlines which are strongly undulate and which are produced at the intersections into nodulose growths. Shell still retaining its natural colour, from olive green to black”. [O1- Measurements (mm.)— length width height 20.0 14.5 10.5 holotype Synonymy— 1923 Cellana hentyi Chapman and Gabriel, Proc. Roy. Soc. Vict., new ser., vol. 36, p. 23, pl. 1, fig. 2. Types—The holotype is in the National Museum of Victoria. Cellana analogia Iredale, 1940 (Pl. 153, figs. 4-6) Range—Lord Howe Island, Roach Islands Remarks—This species is distinguished from howensis, another Lord Howe Island species, mainly in the form of the sculpture, the differ- ences being especially marked when young shells of each are compared. The sculpture in analogia consists of coarse sharply carinated radials that are rendered scabrous to nodulose by dense concentric growth ridges, but in howensis the radials are flattened, have linear interspaces, and weaker concentric sculpture renders the radials only slightly granulose over the earlier growth stages, the ribbing becoming smooth towards the margin in the adult. Description—Shell of moderate size, up to 41 mm. (1% inches) in length, solid, broadly ovate, only moderately elevated, the apex varying from subcentral to the anterior third; margin strongly Plate 153. Figs. 1-3. Cellana howensis Iredale, 1940, Lord Howe Island, 30-32 mm., AWBP. coll. 45412 (one marked X compared with holotype). Figs. 4-6. Cellana analogia Iredale, 1940, Lord Howe Island, 32-39 mm., AWBP. coll. 45413 (one marked X compared with holotype). 174 Subgenus Cellana A. W . B. Powell Patellidae crenulated. Sculpture strong and coarse, con- sisting of numerous carinated radials; primaries more or less alternating with secondaries; linear interspaces deep, and the whole surface ren- dered strongly scabrous to nodulose by dense concentric growth ridges. Young examples are very depressed and have 9 of the primary radials somewhat stronger than the rest. Colour of ex- terior dull-white; interior greyish white, the spatula yellowish to fawn, but mostly completely covered by white callus; margin white-callused, with short dark-brown lines corresponding to the external rib interstices; in some examples these radial colour lines extend intermittently, within the shell substance, almost to the spatula. Measurements (mm.)— length width height 41.0 36.7 20.0 Lord Howe Id. 39.0 33.0 13.0 Lord Howe Id. 37.0 31.0 17.0 holotype OOeD 28.5 10.0 Lord Howe Id. Synonymy— 1940 Cellana analogia Iredale, Aust. Zool., vol. 9 (4), p. 432, pl. 32, figs. 2, 14; pl. 33, figs. 7-9. Types—The holotype and paratypes are in the Australian Museum, Sydney, and there is a series of topotypes in the Powell collection, Auckland. Cellana howensis Iredale, 1940 (PI. 153, figs. 1-3) Range—Lord Howe Island, Ned’s Beach. Remarks—This species is distinguished from the other Lord Howe Island limpet, C. analogia, in the form of the ribbing which consists of broad low radials that are separated by linear inter- spaces. There is also, a radial pattern of dark- brown lines in the rib interstices, as well as vary- ing radial streaks of the same colour. The nearest related species seems to be the Australian tra- moserica. Description—Shell of moderate size, up to 34 mm. (1% inches) in length, rather solid, ovate, elevated, the apex at the anterior fourth; margin finely crenulated. Sculpture consisting of num- erous, flattened, radial ribs of varying width, some rather broad, and all with linear inter- spaces; these radials are further subdivided by one or two shallower radial grooves, and the whole surface is densely and delicately concen- trically crossed by growth lines that render the radials weakly granulose over the early part of the shell; there being a general smoothness of the ribbing towards the margin. Colour of exterior greenish grey to greyish buff, the narrow inter- spaces lined in dark-brown, and some have radial streaks of the same colour; interior yellowish to orange-brown, with the spatula dark reddish brown, usually more or less completely clouded with greyish-white callus; the external pattern shows through strongly except in fully adult examples, which have a rounded callused margin, and in these the external radial lines form short radial dashes corresponding to the external linear interspaces. Measurements (mm.)— length width height 34.0 28.0 13.25. Lord Howe Id. 31.5 26.5 13.50 holotype 31.0 26.0 18.00 Lord Howe Id. 26.0 21.0 11.50 Lord Howe Id. Synonymy— 1940 Cellana howensis Iredale, Aust. Zool., vol. 9 (4), p. 432, pl. 32, figs. 1, 13; pl. 33, figs. 4-6. Types—The holotype and paratypes are in the Australian Museum, Sydney, and there is a series of topotypes in the Powell collection, Auckland. Cellana craticulata (Suter, 1905) (PI. 154, 155; PI 156, fig. 2) Range—Kermadec Islands. Remarks—This is a difficult species to describe in general terms, since it is excessively variable, assuming different shapes, sizes, sculptural de- velopments and colour patterns, presumably in relation to degrees of exposure, and vertical dis- tribution within the tidal belt. This complex was divided by Oliver (1915, pp. 511-514) into 4 species and 2 subspecies, but his interpretation breaks down in practice, as evidenced by Oliver's own qualifying remarks (l.c. p. 511):—*examina- tion of ample material leads one to the conclusion that in the Kermadec group there exist about four species of Cellana in the process of being formed out of a single species, and the young of all are frequently so much alike that a satisfactory dis- position is scarcely possible.” Another significant point is that although three of the named forms have their respective type localities elsewhere in the group than Raoul Island, all six are recorded from that island. Lives on rocks from low to high water. Description—Shell of small to moderately large size, from 21.5 to 50.6 mm. (%-2 inches) in length, [01-728] November 27, 1973 narrowly to broadly ovate, depressed to elevated, with weakly crenulated to strongly corrugated margins, and excessively variable sculpture and colour pattern. The typical form is elongate oval, depressed, and strongly sculptured, the radials basically in the form of distant, broadly rounded corrugations, these and the interspaces, densely overridden by narrow radial cords, which are rendered weakly nodulose by dense concentric growth lines. Coloration; externally, olive with most of the radial folds broadly radially streaked in dark-brown to black; internally, silvery with the radiate external pattern showing through, Plate 154. Figs. 1-9. Cellana craticulata (Suter, 1905), Ker- madec Islands. Figs. 1, 2. Raoul Island (typical form), 26-30 mim., MPIM649. Figs. 3. 4. Raoul Island (prolixa form), 35 min. MPM64S. Pig. 5. Denham Bay, Raoul Island (prolixa form), 29 mim. MPFIM651. Figs. 6, 7. French Rock (scopulina INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana = 175 strongest at the margin; spatula long and_nar- row, dark-sepia, clouded with white callus. The form hedleyi has the radial folds well developed but the outline is more broadly ovate; corrugata is similar but has maximum development of the radial folds; vulcanica is high-conic and coarsely ribbed but without radial folds; scopulina is rounded and high-conic also, but the radial sculpture is not prominent, only occasional young examples have the radial folds, and the general coloration is yellowish to pinkish orange, often with radiate streaks and interstitial dark-brown lines. NILE Gy, os, SS form), 50 mimn., MEI465S. Figs. $8, 9. Raoul Island, (scopulina form), 21-22 min., AWBP coll. (Che ME numbers of this and the following plate refer to Dominion Museum, Wellington specimens in the WR. B. Oliver collection; Oliver's deter- minations in brackets). [01-729] 176 Subgenus Cellana A. W. B. Powell Patellidae Measurements (mm.)— length width height 50.0 42.0 20.8 scopulina form; Oliver, 1915 45.8 37.5 14.2 hedleyi form; Oliver, 1915 42.8 37.0 11.3. corrugata form; Oliver, 1915 39.0 30.5 9.5 hedleyi form; Raoul Id. 32.2 28.0 16.3. vulcanica form; Oliver, 1915 31.4 26.4 7.4 prolixa form; Oliver, 1915 26.5 23.0 7.5 scopulina form; Raoul Id. 25.0 20.0 7.0 holotype of craticulata Synonymy— 1905 Helcioniscus craticulatus Suter, Proc. Malac. Soc., Lond., vol. 6, p. 352, text figs. 1910 Helcioniscus dirus Reeve, (non Reeve, 1855) Iredale, Proc. Malac. Soc., Lond., vol. 9, p. 71. 1910 Helcioniscus craticulatus Suter, Iredale, Proc. Malac. Soc., Lond., vol. 9, p. 72. 1913 Helcioniscus antipodum Smith, (non E. A. Smith, 1874) Suter, Man. N. Z. Moll., p. 79 (in part). 1915 Cellana craticulatus Suter, Oliver, Trans. N. Z. Inst., vol. 47, p. 511. 1915 Cellana craticulatus prolixus Oliver, Trans. N. Z. Inst., vol. 47, p. 512, pl. 9, figs. 1, la. 1915 Cellana hedleyi Oliver, Trans. N. Z. Inst., vol. 47, p. 512, pl. 9, figs. 2, 2a. 1915 Cellana hedleyi corrugata Oliver, Trans. N. Z. Inst., vol. 47, p. 513, pl. 9, figs. 3, 3a. 1915 Cellana vulcanicus Oliver, Trans. N. Z. Inst., vol. 47, p. 513, pl. 9, figs. 4, da. 1915 Cellana scopulinus Oliver, Trans. N. Z. Inst., vol. 47, p. 514, pl. 9, figs. 5, 5a. Plate 155. Figs. 1-11. Cellana craticulata (Suter, 1905), Kermadec Islands (continued). Figs. 1-3. Meyer Island (vul- canica form), 25-35 mm. (Fig. 3, compared with holotype), MP14664 & MPI4665. Figs. 4-6 Raoul Island (hedleyi form), 32-46 mm., MF14659. Figs. 7, 8. Coral Bay, Raoul Island (hedleyi corrugata form), 35 mim., MPF14653. Figs. 9-11. Macaulay Island (hedleyi corrugata form), 35-44 mi., MEF3082. [01-730] November 27, 1973 Plate 156. Fig. 1. Cellana radians (Ginelin), New Zealand, Ti Point, Hauraki Gulf. Radula. Fig. 2. Cellana craticulata (Suter), Macaulay Island, Kermadecs. Radula. Types—The holotype of craticulatus is in the Suter collection, New Zealand Geological Survey, Wellington, and the Oliver collection is in the Dominion Museum, Wellington. Records—KERMADEC ISLANDS: Raoul or Sunday Island (holotype); (Auck. Mus.; AWBP coll.); Meyer Islet (AWBP coll.); Macaulay Island (Oliver, 1915); French Rock (Oliver, 1915; AWBP coll.). Cellana denticulata (Martyn, 1784) ! (Pl. 70, figs. 7, 8; Pl. 157, figs. 5, 6; Pl. 163, fig. 2) Range—New Zealand: Three Kings Islands, North Island, and northern part of South Island. Remarks—This shell is readily distinguished by its prominent brown scaly ribs, netted with brown in the interstices, and by its internal coloration, the spatula being cream to orange-brown, and the rest of the interior rayed and netted in dark- brown upon a bluish grey ground. This is the dominant limpet of the Cook Strait area, and from there it extends southward to at least Kaikoura. In its northern range, up the North Island east coast, it reaches the Three Kings Islands, but is not generally distributed in the north. These northern isolated small colonies are situated on certain jutting points and off shore islands, which evidently are catchments for larvae transported by coastwise currents. Description—Shell of moderate to large size, up to 74 mm. (almost 3 inches) in length, solid, elevated; regularly, closely and strongly radially ' This name and others of Martyn, 1784, relevant to the New Zealand fauna, were validated by the International Com- mission of Zoological Nomenclature, in Opinion 479 (1957). INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 177 ribbed, the whole surface crossed by numerous lamellose concentric ridges that thicken into granular scales on the radials. Colour of exterior greyish, with the radials and a netted interstitial pattern in dark brown; interior brownish with the external pattern showing through strongly in brown or purplish brown. The well defined spatula varies from cream to orange-brown, and there is sometimes a bluish white area between there and the brown blotched margin. Measurements (mm.)— length width height 74.0 63.0 37.0 Karewa Island 73.0 61.0 44.0 Karewa Island TAs 60.0 29.0 Mt. Maunganui 52.0 43.0 19.0 Island Bay 43.5 35.0 15.0 Island Bay Plate 157. Figs. 1-4. Cellana ornata (Dillwyn, 1817), New Zealand. Fig. 1. Motutara, West Coast, Auckland, 21 min.. AWBP. coll. Fig. 2. Charleston, West Coast, South Island (surf-beaten form), 42 mim., AWBP coll. 310. Fig. 3. Bluff, Southland, 42 mm., AWBP coll. 52467. Fig. 4. Campbell's Bay, Auckland, 34 mm., AWBP. coll. 26270. Figs. 5, 6. Cellana denticulata (Martyn, 1784), New Zealand, Island Bay. Wel- lington, 45-48 min., AWBP coll. 292. [01-731] 17S Subgenus Cellana Radula—Formula (3) + 1 + (1+0+1) + 1 + (3). Radula very similar to that of radians, with both the pair of functional centrals and the pair of laterals with long lanceolate cusps set tangen- tially to the shank, and the lower cutting edge of the lateral is indented in two places. The non- functional fused marginal plates have three short parallel ridges at the base as in radians and flava. Synonymy— 1784 Patella denticulata Martyn, Univ. Conch., vol. 2, fig. 65. Name validated by I. C. Z. N., opinion 479 (1957). 1855 Patella imbricata Reeve, Conch. Iconica, pl. 32, figs. 93 a, b. 1880 Patella reevei Hutton, Man. N. Z. Moll. p. 108, nom. nov. pro P. imbricata Reeve, 1855, non Turton, 1802. IS9L Helcioniscus denticulatus: Pilsbry, Man. Conch., vol. 13, p. 138, pl. 68, figs. 23, 24; pl. 21, figs. 49, 50. 1913 Helcioniscus denticulatus: Suter, Man. N. Z. Moll, p. 80, pl. 7, fig. 10. 1957 Patella denticulata Martyn; validation of name, I. C. Z. M. opinion 479, p. 369. Records—NEW ZEALAND: Three Kings Islands, Great Island (AWBP); North Island; Cape Maria van Diemen (AWBP); Karewa Island, Bay of Plenty (Auck. Mus.); Mt. Maunganui (AWBP coll.); Island Bay, Wellington (AWBP coll.); South Island; Goose Bay, Kaikoura (AWBP coll.). Cellana flava (Hutton, 1873) (Pl. 70, figs. 5, 6; Pls. 158, 159) Range—New Zealand, east coast of both North and South Islands, from East Cape to Motanau Island, north Canterbury. Remarks—This_ bright yellowish to orange limpet, long considered to be only a subspecies of radians, merits specific separation from that species, not only on account of its distinctive Plate 158. Cellana flava (Ilutton, 1873), New Zealand. Fig. . East Cape. North Island, 41 mim., AWBP coll. 52732. Figs. {. Limestone Point, Marlborough, South Island, 1S-51 min. Note the dark rays found occasionally in some young ex- amples; AWBP coll. 6872: Figs. 3, 4). | 2. ( [01-732] A. W. B. Powell Patellidae coloration but also, by virtue of its simple yet relatively constant form of sculpture, short stubby cephalic tentacles, and peculiar vestigial central radular plate. The species has its centre of distribution along the Kaikoura-Amuri Bluff coast of Marlborough Where it occurs in great numbers on white lime- stone in the mid- to low-tidal zone. Like denticu- lata, its extra-limital occurrences are sporadic, and almost always on prominences of the coast- line. The pale coloration possibly resulted from long association with a white limestone sub- stratum, but if so, it cannot be a direct response, for bright-orange examples are just as likely to be found living on dark rock, notably at Whaka- take, near Castle Point, in the North Island. Description—Shell of moderate size to rather large, up to 66 mm. (over 2% inches) in length, solid, elevated, with the apex varying from sub- central to about the anterior third. Sculpture simple and rather constant, consisting of from 19 to 25 strong, rounded, primary radials, with an occasional! much weaker intermediate that devel- ops only towards the margin. Colour, both ex- ternally and internally, pale-yellow to bright- orange. In senile examples the exterior is usually eroded to dull grey or whitish, and the interior is whitish also, except for the large spatula, which is invariably yellowish to orange. Occasional Plate 159. Cellana flava (Hutton), New Zealand, East Cape. Radula. November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 179 young shells have several irregularly disposed brown radial stripes developed only towards the margin. Radula—Formula 3 + 1 + (1+(1)+) + 1 + 3; very similar to the radula of radians, even to the ap- pearance of a vertical alternation of long and short marginals, but the vestigial central plate is different, being foliated, open above, and with the front edge scalloped into five cusp-like lobes (East Cape example). Measurements (mm.)— length width height 66.0 55.0 27.0 Karaka Point 58.0 48.5 28.0 Limestone Point 47.5 37.5 19.5 Limestone Point 35.5 29.5 14.5 Limestone Point Synonymy— 1873 Patella flava Hutton, Cat. N. Z. Moll., p. 44. 1891 Helcioniscus flavus: Pilsbry, Man. Conch., vol. 13, p. 142. 1913 Helcioniscus radians flavus: Suter, Man. N. Z. Moll., p. 54; Atlas (1915), pl. 7, fig. 18. Types—The type is in the Dominion Museum, Wellington. Records—NEW ZEALAND: North Island; Horoera, East Cape; Gisborne; Tolaga Bay; Whakatake; Castlepoint, East Wairarapa. South Island: Karaka Bay, Queen Charlotte Sound; Limestone Point, S. of Clarence River, Marlborough (all AWBP. coll.); Kaikoura, Amuri Bluff and Motanau Island (Suter, 1913). fe JF fae it Plate 160. Figs. 1-9. Cellana radians (Gmelin, 1791), New Zealand, 23-65 mim. Fig. LL. earlii pattern, Motutara, West Coast, Auckland, AWBP coll. 252. Figs. 2-5. Mount Maun- gamui, Bay of Plenty, AWBP coll. 52469, 293 & 299. Figs. Cellana ornata (Dillwyn, 1817) (PI. 70, figs. 12, 13; Pls. 157, 163) Range—New Zealand: North, South and Stewart Islands. Remarks—In its young non-eroded state this species has a most attractive colour pattern, the primary ribs being lilac-grey, and the interme- diate areas purplish brown to black, with the nodes picked out in white. In large-sized examples the exterior is almost invariably eroded and little of the basic colour pattern remains. Also, in coastal areas subject to rigorous wave action, the profile is low, and the shape varies from elongate-ovate to broadly-ovate. This species. is common on rock faces in the upper tidal zone. Description—Shell small to moderate sized, up to 48 mm. (almost 2 inches) in length, but usually between 24 and 28 mm., solid, normally high-conical, with the apex at about the anterior third. Sculpture consisting of eleven strong, pri- mary, radial ribs, each interspace with a central nodulose secondary radial, flanked on each side by a pair of much weaker radials; the whole crossed by dense concentric threads, that thicken like knots, wherever they surmount the primary and secondary radials. Colour of ex- terior: the primary ribs pale lilac-grey, the inter- spaces dark purplish brown to almost black, 6, 7. Four miles south of Clarence River, Marlborough, AWBP coll. 3783. Figs. $8, 9. Motuihi Island, Hauraki Gulf, Auck- land, AWBP coll. 18600. [01-733] 180) Subgenus Cellana A. W. B. Powell Patellidae with the nodes on the secondary radials picked out in white. Color of interior: with broad, dark purplish brown rays and _ silvery intermediate narrow rays; spatula dark chestnut-brown to almost black, often partly clouded with greyish callus. Radula—Formula (3) + 1 + (1+0+1) + 1 + (3); similar to the radula of radians, the paired cen- trals each with a long lanceolate cusp, set tan- gentially to the shank, and the laterals similar but heavier, with the lower edge indented to form two denticles; the three marginals are fused into a small irregularly-shaped, semitransparent plate, without cusps. Measurements (mm.)—(all A.W.B. Powell col- lection). length width height 48.0 44.5 21.0 Motutara 44.0 34.0 11.0 Charleston 42.5 35.0 21.0 — Bluff 42.0 35.0 19.0 Mt. Maunganui 28.0 92:0 13.0 Motutara 22.5 18.5 7.0 Motutara Synonymy— 1817 Patella ornata Dillwyn, Cat. Rec. Shells, vol. 2, p. 1029; based upon Martini-Chemnitz, Conch. Cab., vol. 11, p. 180, pl. 197, figs. 1914, 1915. 1841 Patella nodosa Hombron & Jacquinot, Ann. Sci. Nat., vol. 2 (16), p. 191. 1846 Patella luctuosa Gould, Proc. Boston Soc. Nat. Hist., vol. 2, p. 150. 1855 Patella’ margaritaria Reeve, Conch. Iconica, pl. 28, figs. 74 a, b. 1883 Patella ‘denticulata’ (error for ornata); Hutton, Trans. N. Z. Inst., vol. 15, p. 128, pl. 16, fig. B (radula). 1891 Helcioniscus ornatus: Pilsbry, Man. Conch., vol. 13, p. 137, pl. 68, figs. 14-19: pl. 19, figs. 39, 40. 1913 Helcioniscus ornatus: Suter, Man. N. Z. Moll., p. 80; Atlas (1915), pl. 7, fig. 11. 1913 Helcioniscus ornatus inconspicuus: (non Gray, 1843), Suter, Man. N. Z. Moll., p. 81; Atlas (1915), pl. 7, fig. 12. Records—NEW ZEALAND: North Island; Cape Maria van Diemen (Auck. Mus.); Reotahi, Whangarei Heads; Motuihi Island, Auckland; Campbell's Bay, Auckland; Motutara, West Coast, Auckland; Mt. Maunganui, Bay of Plenty; Napier. South Island: 4 mi. S. of Clarence River, Marlborough; Lyt- telton; Cape Foulwind; Charleston; Oamaru; Kartigi Beach, north Otago (all AWBP coll.); Taieri Beach, Otago (Auck. Mus.); Solander Island, Foveaux Strait (Auck. Mus.). Stewart Island; Herekopare Island (both AWBP coll.). Cellana radians (Gmelin, 1791) (Pl. 70, figs. 1-4; Pls. 156, 160, 161) Range—New Zealand: North, South and Stewart Islands. Remarks—This is the most common of the New Zealand limpets and the most variable, not only in shape and sculpture, but also in colour pattern. The many forms of the species are outlined in the following formal description. In general, north- ern shells, which are the typical form, have the primary ribs coloured brown, and there is often a connecting pattern of transverse streaks (the earlii pattern). Most southern shells, on the other hand, have the sculpture finer, more even, the external markings indistinct, and internally there is a greenish silvery to golden lustre. This, the perana form, is the dominant one at Stewart Island and the southern part of the South Island, but it is known to occur also on the west coast of the North Island at Whitecliffs, north Taranaki, and also at the Three Kings Islands. On the other hand, at Cape Foulwind on west coast of the South Island both the perana and earlii forms occur together. The earlii pattern, which is more common in northern shells, is essentially a juve- nile one, and seldom persists into the fully adult, without resolving into radial streaks, more or less confined to the primary radials. Thomson (1919), in his paper on polymorphism in Cellana radians was of the opinion that colour pattern changes in the fully adult of this species were due to external erosion, accompanied by a compensating internal build-up of callus. In such senile examples only deep-seated colour, associated with the primary radials still persists. Helcioniscus radians mestayerae Suter, 1906, is not a New Zealand shell, despite the cited locality, Stewart Island, but is based upon a wrongly labelled specimen of the Indo-Pacific Cellana testudinaria Linnaeus, 1758. Description—Shell of medium to moderately large size, up to 65.5 mm. (2% inches) in length, polymorphic, extremely variable in shape, alti- tude and colour pattern. Typical form ovate, depressed, with the apex at about the anterior fourth; sculptured with 20 to 25 narrow, slightly raised primary radials, and a varying number of very weak radial threads in the interspaces, but often, the latter are subobsolete; the whole sur- face crossed by weak, but exceedingly dense, concentric lirations. Colour of exterior greyish buff, with a reddish brown pattern of interrupted radial lines and transverse streaks; interior yel- lowish, with the external brown markings show- ing through strongly; spatula ill-defined, fawn to chestnut-brown. In the earlii form the trans- verse streaks are dominant, and join up in a con- centric anastomosing pattern. In the decora form the pattern is restricted to radial lines; and in the perana form the sculpture is fine and more regu- lar, the external coloration of dark, continuous [01-734] November 27, 1973 or intermittent, radial lines, and internally it is greenish silvery to golden, sometimes partly clouded to fully obscured by a white callus. Radula—Formula 3 + 1 + (1+0+1) + 1 + 3. There is a pair of strong centrals, each with a long lanceolate cusp, set tangentially to the shank, and in between these two teeth is a small, narrow, vestigial median plate; the pair of laterals are similar but have a broadly triangular base, and the lower cutting edge of the cusp is indented to form two denticles; all three marginals are pres- ent but they are very thin and semitransparent; only the inner one bears a slight cusp, and below these, joined by a thin membrane are three shorter INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 181 narrow plates, the effect being of long and short marginals in a vertical alternation. Measurements (mm.)—(As=typical form; B= earlii form; C=perana form. All from the A. W. B. Powell coll’n. length width height 65.5 55.0 24.0 (A) Mt. Maunganui 64.5 54.5 26.0 (C) Herekopare Island 62.0 50.0 30.0 (C) Herekopare Island 59.0 50.5 16.0 (A) Mt. Maunganui 57.0 48.0 20.0 (C) Herekopare Island 46.0 37.5 12.0 (A) Motuihi Island 43.5 35:5 17.0 (C) Cape Foulwind 41.5 34.0 20.5. (C) Cape Foulwind 35.5 28.0 7.75 (B) Cape Foulwind 28.0 29.5 6.0 (A) Little Barrier Id. 18.5 15.0 3.0 (B) Little Barrier Id. Plate 161. Figs. 1-7. Cellana radians (Gmelin, 1791) (perana form). Figs. 1, 2. Herekopare Island, Stewart Island, 57 mim., AWBP coll. 26449. Figs. 3, 4. Lyttelton, South Island, 41-4 no., AWBP coll. 301. Figs. 5-7. North West Landing. Great Island. Three Kings Islands. New Zealand. 25-35 mm. AWBP coll. 52657. [01-735] 182. Subgenus Cellana A. W. B. Powell Patellidae Synonymy— 1791 Patella radians Gmelin, Syst. Nat., ed. 13, p. 3720; based upon Martini-Chemnitz, Conch. Cab., vol. 10, pl. 168, fig. 1618. 1830 Patella argyropsis Lesson, Voy. Coquille, Zool., vol. 2, p. 419. 1830 Patella pholidota Lesson, Voy. Coquille, Zool., vol. 2, p. 420. 1834 Patella argentea Quoy and Gaimard, Voy. Astrolabe, Zool., vol. 3, p. 345, pl. 70, figs. 16, 17. 1841 Patella radiatilis Hombron and Jacquinot, Ann. des Sci. Nat., vol. 16, p. 191. ?1848 Patella orichalcea Philippi, Zeitschr. f. Malak., p. 163. 1549 Patella decora Philippi, Zeitschr. f. Malak., p. 162. 1854 Patella decora Philippi, Reeve, Conch. Iconica, pl. 15, figs. 33 a-c. 1855 Patella earlii Reeve, Conch. Iconica, pl. 27, figs. 71 a, b. 1873 Patella flexuosa (non Quoy and Gaimard, 1834), Hut- ton, Cat. Mar. Moll. N. Z., p. 45. 1874 Patella antipodum E. A. Smith, Voy. Erub. & Terr. Moll, p. 4, pl. 1, fig. 25. 1882 Patella olivacea Hutton, N. Z. Journ. Sci., vol. 1, p. 69. 1891 Helcioniscus radians Gmelin, Pilsbry, Man. Conch., vol. 13, p. 139, pl. 23, figs. 4, 6, 7, 8; pl. 69, figs. 25-31, 34-37 (non figs. 32, 33, 38 & 39). 1913 Helcioniscus radians Gmelin, Suter, Man. N. Z. Moll, p. $1, pl. 7, fig. 13. 1913 Helcioniscus radians argenteus Q. and G., Suter, Man. N. Z. Moll., p. 82. 1913 Helcioniscus radians decorus Philippi, Suter, Man. N. Z. Moll., p. 82. 1913 Helcioniscus radians earlii Reeve, Suter, Man. N. Z. Moll., p. 83. 1913 Helcioniscus radians olivaceus Hutton, Suter, Man. N. Z. Moll, p. $4. 1915 Cellana radians perana Iredale, Trans. N. Z. Inst., vol. 47, p. 432; nom. nov. pro Patella olivacea Hutton, 1882; non Anton, 1839. Plate 162. Cellana_ stellifera (Gmelin, 1791), New Zealand. Figs. 1, 2. Island Bay, Wellington, 33-50 mm., AWBP. coll. 258. Figs. 3, 4. Rocks Road, Nelson, 29-33 mm., AWBP. coll. 45478. 1919 Cellana radians Gmelin, Thomson, N. Z. Journ. Sci. Tech., vol. 2, pp. 264-267 (polymorphism). 1923 Helcioniscus radians Gmelin, Eales, Brit. Antarct. (‘Terra Nova’) Exped., 1910, Moll., pt. 5, pp. 3-6, text. fig. 2 (radula). Records—NEW ZEALAND (typical form): Three Kings Islands. Great Island (F. Climo. 1970); North Island: Cape Maria van Diemen (Auck. Mus.); Busby Head, Whangarei Heads; Little Barrier Island; Motuihi Island, Auckland; Motu- tara, west coast, Auckland; Mt. Maunganui; Gisborne; Tolaga Bay; Island Bay, Wellington. South Island; Cape Foulwind (all AWBP coll.); Lyttelton (AM.); Dowling Bay, Dunedin (Auck. Mus.). Stewart Island; The Neck, Patterson Inlet (AWBP coll.). (perana form): North Island: White Cliffs, north Taranaki. South Island; Goose Bay, Kaikoura; Cape Foulwind; Wainui, Akaroa; Lyttelton; Charleston; Purakanui, Otago; Timaru, Otago; St. Clair, Dunedin; Ocean Beach, Bluff. Stewart Island; Herekopare Island (all AWBP coll.). Cellana stellifera (Gmelin, 1791) (Pl. 70, figs. 9-11; Pls. 162, 163) Range—New Zealand; North, South and Stew- art Islands. Remarks—This species lives at and just below low tide on smooth rock faces in clean water situations but is not generally common. It is easily recognised by its reddish brown external colour, bluish silvery interior, and usual presence of an apical star in paler colour. It is more abundant in the northern part of its range. Description—Shell of moderate size, 30 to 71 mm. (1% to 2% inches) in length, broadly ovate and of low to moderate height, with the apex at about the anterior third, sculptured with numer- ous low rounded radial ridges, crossed uy dense delicate concentric growth lamellae; margin weakly scalloped. Colour, externally dark red- dish brown, mostly showing a white or pale yel- lowish star at the apex, this often persisting to the adult stage, and occasionally with long rays extending from the points of the star right to the margin; interior bluish or purplish grey with a silvery sheen, the star pattern usually showing through; spatula ill-defined, a chestnut smeer often clouded by a whitish callus. Measurements (mm.)— length width height 71.0 58.0 19.0 Whangarei Heads 57.5. 47.5 19.0 Whangarei Heads 45.5 37.0 14.0 Whangarei Heads 33.0 26.5 10.5 Rocks Road, Nelson Synonymy— 1791 Patella stellifera Gmelin, Syst. Nat. ed. 13, p. 3719, based upon Martini-Chemnitz, Conch. Cab., vol. 10, pl. 168, fig. 1617. [01-736] November 27, 1973 1834 Patella stellularia Quoy and Gaimard, Voy. Astrolabe, Zool., vol. 3, p. 347, pl. 70, figs. 18-20. 1855 Patella stellularia Q. and G. Reeve, Conch. Iconica, pl. 33, figs. 96 a, b. 1891 Helcioniscus stellifera Q. and G., Pilsbry, Man. Conch., vol. 13, p. 141, pl. 70, figs. 43-45. 1905 Helcioniscus stelliferus phymatius Suter, Proc. Malac. Soc., Lond., vol. 6, p. 350, text fig. 1913 Helcioniscus stelliferus Q. and G., Suter, Man. N. Z. Moll., p. 86; Atlas (1915), pl. 7, fig. 21. 1915 Cellana_ stellifera: Iredale, Trans. N. Z. Inst., vol. 47, p. 432. Types—The Martini-Chemnitz specimens are possibly in the University Museum, Copenhagen. The cited type locality “Friendly Islands” is erroneous. Records—NEW ZEALAND: North Island; Cape Maria van Diemen (Auck. Mus.); Busby Head, Whangarei Heads; Little Barrier Island; Kawau Island; Mt. Maunganui; Island Bay, Wellington. South Island; Rocks Road, Nelson (all AWBP coll.); New Brighton (Suter, 1913). Stewart Island; Euchre Creek (AWBP coll.). (Suter’s Campbell Island record is based erroneously upon Patinigera terroris (Filhol, 1880)). INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 183 Plate 163. Radulae of New Zealand Cellana. Fig. 1. Cellana stellifera (Gmelin), Ti Point, Hauraki Gulf, Auckland. Fig. 2. Cellana denticulata (Martyn), Makara, Wellington. Fig. 3. Cellana ornata (Dillwyn), East Cape, North Island. In figs. 2 and 3 the centrals and laterals are shown in semi-profile. Key to the subspecies of Cellana strigilis 1. Nucleus at anterior third to seventh of shell A. Shell held to light showing pale spots and shapes (ocellate) a. Outline broadly ovate External colour bluish white, rayed and blotched with light-brown; internal pattern dark-rayed, interrupted at margin 2.0.0.0... eeeeeeeeeeeeeeees redimiculum External colour almost completely clouded sooty-grey to dark-brown; internal pattern dark-rayed, confluent at Margin ........... eee eeeeeteeeeeeeeeteeeeeeees strigilis b. Outline narrowly ovate External colour almost completely clouded with olive-brown; internal pattern dark-rayed, connected at Marin <..6.ssccecccierssilaecalsaiecsveseens flemingi B. Shell held to light showing dense pattern of irregular narrow radials External colour greenish grey, with dense pattern of brown radials; internal pattern dark-rayed, interrupted at margin Outline narrowly ovate ..........ce Outline broadly ovate ............c:cceeee sefos sacs bone ice tet ted atinar ase tetenete ae decease eae bollonsi Fesnocagtt cite seca at ater shea Gna tnevaioutieanenicecs chathamensis 2. Nucleus at anterior 10th to 27th of shell; shell held to light showing heavy radial streaks; outline narrowly ovate; external colour bluish white, rayed and heavily blotched; internal pattern dark-rayed, interrupted at margin Cellana strigilis (Hombron and Jacquinot, 1841) Range—South Island, Stewart Island, Chatham Islands, and southern islands of New Zealand. Remarks—The genus Cellana is typically warm- water Indo-Pacific distribution, so it is remark- able to find the genus extending to as far south as Campbell Island, 52° 30’S. These occurrences are probably relict from former warmer geologi- Fe em tn rl re Ore Teen ee ee ET ems Fe oliveri cal times, for all the islands concerned stand upon the extensive submarine platform surround- ing New Zealand (See Plate 99). Subspecies have developed in isolation, and although they are all closely allied, recognisable differences are apparent in the shells from these segregated populations. A key to these subspecies follows: [01-737] 154. Subgenus Cellana Cellana strigilis subspecies strigilis (Hombron and Jacquinot, 1841) (Pl. 70, figs. 14, 15; Pls. 164, 168) Range—Auckland and Campbell _ Islands, southern New Zealand. Remarks—The southern islands’ strigilis and the mainland redimiculum are closely allied, but always easily distinguished by the fact that the former is dark, with the interior sooty-grey, having an underlying densely mottled pattern in dark-brown, which forms an approximately con- tinuous dark margin to the shell. On the other hand, redimiculum is yellowish brown, with dark A. W. B. Powell Patellidae reddish brown radials that do not anastomose at the margin. Description—Shell large, up to 80 mm. (3% inches) in length, solid, broadly-ovate, elevated, with the apex varying between the anterior third and sixth. Sculpture consisting of 20 to 25 strong, rounded, radial ribs, mostly with a weaker radial in each interspace; the whole surface crossed by dense, weak, concentric growth lines. Colour of exterior dark brown or greenish, to almost black, with a few spots and streaks of yellowish-white, much more prominent when the shell is held- to the light; interior purplish-brown to sooty-grey, clouded and indistinctly rayed with dark brown; Plate 164. Cellana. strigilis (Hombron & Jacquinot, S41). Figs. 1, 2. Perseverance Harbour, Campbell Island, 78 mm., AWBP. coll. 26165. Fig. 3. Under South Col, Campbell Is- land, 37 mm., AWBP coll. 42168. Fig. 4. Garden Cove, Camp- bell Island, 46 mim., AWBP coll. 42169. [01-738] November 27, 1973 spatula light yellowish-brown, irregularly mar- gined in dark grey. In non eroded young shells the ground colour varies from russet-brown through greenish grey to almost black, with the interstitial odd spots and splashes pale bluish. Measurements (mm.)— length width height 80.0 68.0 39.0 Campbell Island 77.5 66.5 30.0 Campbell Island 70.0 60.0 34.0 Shoal Pt., Campbell Id. 65.0 52.0 46.0 Shoal Pt., Campbell Id. 57.0 42.5 17.0 Garden Cove, Campbell Id. 38.0 31.0 12.5 Garden Cove, Campbell Id. Synonymy— 1841 Patella strigilis Hombron and Jacquinot, Ann. Sci. Nat., vol. 2, pt. 16, p. 190. 1846 Patella illuminata Gould, Proc. Boston Soc. Nat. Hist., vol. 2, p. 149. 1891 Helcioniscus strigilis H. and J., Pilsbry, Man. Conch., vol. 13, p. 137. 1891 Helcioniscus illuminata Gould, Pilsbry, Man. Conch., vol. 13, p. 142, pl. 70, figs. 40-42. 1913 Helcioniscus strigilis (in part): Suter, Man. N. Z. Moll., p. 87. 1924 Cellana radians Gmelin, Odhner, N. Z. Moll., Pap. Mor- tensen Pacific Exped., p. 11 (non Gmelin, 1791). 1927 Nacella strigilis H. and J. Finlay, Trans. N. Z. Inst., vol. 57, p. 387. 1955 Cellana strigilis strigilis H. and J. Powell, D. S. I. R., Cape Exped. Ser., Bull. no. 15, p. 70. Types—The type of strigilis is in the Muséum National d’ Histoire Naturelle, Paris, and that of illuminata in the United States National Mu- seum, Washington. Records—Southern islands of New Zealand: AUCKLAND ISLANDS (type): Musgrave Peninsula; Tagua Bay, Carnley Harbour; Crozier Point; Waterfall Inlet; Rose Island, Port Ross, and Enderby Island (NZGS); Carnley Harbour; Han- field Inlet (both AWBP. coll.). CAMPBELL ISLAND, Perse- verance Harbour (NZGS; AWBP. coll.); Shoal Point (Auck. Mus.); Monument Harbour (Cape Exped., 1945). Cellana strigilis subspecies bollonsi Powell, 1955 (PI. 165, figs. 1,2) Range—Antipodes Islands, southern New Zea- land. Remarks—This subspecies is easily recognised by its dense pattern of interstitial brown lines and streaks on a greenish grey ground. Occa- sionally the subspecies chathamensis has a simi- lar pattern in juvenile shells, but it never persists into the adult stage, as it does invariably in bollonsi. Description- -Shell of moderate size to relatively large, up to 70.5 mm. (2% inches) in length, solid, nerrowly ovate, depressed to moderately eleva- INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana = 185 ted, with the apex ‘varying between the anterior fifth and seventh. Sculpture developing from scarcely raised radial folds in juveniles to from 20 to 24 narrowly rounded, sharply raised ribs in the adult. Colour of exterior greenish-grey, the radials marked out in light-brown to reddish- brown, plus a dense overall pattern, in these same colours, in the form of interstitial meander- ing radial lines and streaks; interior metallic dull blue-grey, with reddish-brown external pattern showing through; spatula buff to pale brown. Measurements (mm.)— length width height 70.5 58.0 28.0 — Antipodes Ids. 62.0 49.0 21.0 Antipodes Ids. 48.5 36.0 15.0 — Antipodes Ids. 48.0 34.8 14.5. Antipodes Ids. 42.0 32.0 12.0 Antipodes Ids. Synonymy— 1955 Cellana strigilis bollonsi Powell, Dept. Sci. and Indust. Res., Cape Exped. Ser., Bull. no. 15, p. 73, pl. 5, figs. 51-53. Types—The holotype and paratypes are in the Dominion Museum, Wellington. Records—ANTIPODES ISLANDS (holotype and paratypes); (AWBP coll.). Plate 165. Figs. 1, 2. Cellana_ strigilis subspecies bollonsi Powell, 1955, Antipodes Islands, 48-62 mm., AWBP. coll. 26197, 258420. Figs. 3-5 Cellana_ strigilis subspecies oliveri Powell, 1955, Bounty Islands, 35-47 mm., AWBP coll. 26198 (01-739] 186 Subgenus Cellana A. W. B. Powell Patellidae Cellana strigilis subspecies chathamensis (Pilsbry, 1891) (Pl. 167; Pl. 168, fig. 1) Range—Chatham Islands, New Zealand. Remarks—This subspecies resembles redimicu- lum in its simple radiate reddish brown radials, not coalescent at the margin, but the shape is more roundly arched, and the ocellate pattern is not in evidence. On the other hand some juve- niles have a dense meandering pattern, remini- scent of the bollonsi pattern. Description—Shell of moderate to large size, up to 73.5 mm. (2% inches), in length, solid, ovate, elevated, and roundly arched in profile, with the apex at about the anterior third. Sculpture con- sisting of from 21 to 25 moderately strong, NORTH ID _—GHATHAM IDS) ek. ta)eP TIPODES 105] 50° 09 5004000 1000—FATH Plate 166. Distribution of Cellana strigilis and its subspecies. An example of an otherwise warm-water genus, surviving cooling temperatures in southern New Zealand, and repre- sented as relict subspecies in the isolated southern islands, once part of a greater New Zealand land mass. A- Cellana. strigilis redimiculum (Reeve), South Island and Stewart Island. Note northern limit near top of South Island east coast. B- Cellana strigilis flemingi Powell, Snares Islands. C- Cellana strigilis strigilis (Hombron & Jacquinot), Auck- land Islands and Campbell Island. D- Cellana strigilis bollonsi Powell, Antipodes Islands. K- Cellana strigilis oliveri Powell, Bounty: Islands. F- Cellana strigilis chathamensis (Pilsbry), Chatham Islands. (Chart adapted from Fleming, 1951, N. Z. Science Review, 9 (10), p. 167). rounded, radial ribs, with a weaker one in most interspaces; the whole surface crowded with con- centric growth lirations, weak over the early stages of the shell but stronger towards the mar- gin, where they become slightly knotted across the radials. Colour of exterior pale yellowish- brown to greyish lilac, the radials lined in red- dish brown; internally, silvery to yellowish brown, with regular reddish brown radial lines, corres- ponding to the external ribbing, their terminal points not coalescent at the margin; spatula large, fawn to orange-brown. Juvenile shells pale yellowish to almost black, with the pattern vary- ing from a few radial lines to a dense coverage of meandering lines and streaks. Measurements (mm.)— length width height 73.5 56.5 35.0 Chatham Island 70.0 55.0 31.0 Chatham Island 62.0 49.0 23.5 Chatham Island 57.5 47.5 27.0 Waitangi, Chathams 56.5 45.5 20.0 Waitangi, Chathams 39.5 30.0 11.0 Pitt Id., Chathams Synonymy— 1891 Acmaea_ chathamensis Pilsbry, Man. Conch., vol. 13, p. 56, pl. 35, figs. 43-46. 1933 Cellana chathamensis: Powell, Rec. Auck. Inst. Mus., vol. 1 (4), p. 196, pl. 36, figs. 1-4. 1955 Cellana_ strigilis chathamensis: Powell Dept. Sci. and Indust. Res., Cape Exped. Ser., Bull. no. 15, p. 73. Types—The type material is in the Academy of Natural Sciences of Philadelphia. Records—NEW ZEALAND: CHATHAM ISLANDS (type): Waitangi (AWBP coll); Wharekauri (AWBP. coll.); Tioriori (Auck. Mus.); Waihere Bay, Pitt Island (Auck Mus.). Cellana strigilis subspecies flemingi Powell, 1955 (Pl. 168, fig. 3; pl. 169, figs. 5-7) Range—Snares Islands, southern New Zealand. Remarks—This subspecies is more closely allied to typical strigilis than it is to redimiculum, from both of which it differs in its consistently more narrowly oval shape, high arched profile, and anterior position of the nucleus. Description—Shell of small to medium size, up to 53 mm. (2% inches) in length, narrowly ovate, with the apex varying between the anterior fourth to fifth in adults, but one eighth or less in juveniles. Anterior slope straight, but posterior slope prominently arched, and flattened on top [01-740] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 187 Plate 167. Figs. 1-4. Cellana strigilis subspecies chathamen- sis (Pilsbry, 1891). Waitangi, Chatham Islands, 21-71 mm., AWBP coll. 8756, STSS. for about one third of the length. Sculpture con- sisting of about 25 narrowly rounded primary radials, and a few very weak intermediates. Young shells have the radials as scarcely raised folds, crossed by dense concentric growth lines. Colour of exterior almost uniformly olive-brown, except for the nuclear area to about 15 mm., which is dark-brown, with a light bluish ocellate pattern in the rib interstices; interior metallic dull smoky-grey, with a dark brown internal rib pattern showing through the glaze, and coalesc- ing at the margin, in adults, to form an almost continuous border; spatula buff, tinged posteriorly with pale reddish brown. Measurements (mm.)— length width height 53.00 41.5 22.00 holotype 47.00 35.5 18.50 paratype 28.75 21.0 7.00 paratype 21.25 15.0 4.25. paratype 17.75 12.5 3.40 paratype Synonymy— 1955 Cellana_ strigilis flemingi Powell, Cape Exped. Ser., Bull. no. 15, p. 72, pl. 5, figs. 45-47 Types—The holotype and paratypes are in the New Zealand Geological Survey, Wellington. Records—SNARES ISLANDS: boat harbour, on intertidal rocks. Cellana strigilis subspecies oliveri Powell, 1955 (Pl. 165, figs. 3-5) Range—Bounty Islands, southern New Zealand. Remarks—This subspecies is easily recognised by its narrowly ovate and depressed shape, with the apex at, or near to, the anterior end, and a bold pattern of radial streaks and blotches. Description—Shell of small to medium size, up to 57.5 mm. (2% inches) in length, solid, rather narrowly ovate and depressed, the apex near to the anterior margin at all stages of growth. From [01-741] 188 Subgenus Cellana A. W. B. Powell Patellidae Plate 168. Radulae of New Zealand Cellana. Fig. 1. Cellana strigilis (HMombron & Jacquinot), Campbell Island. Fig. 2. Cellana_ strigilis chathamensis (Pilsbry), Wharekauri, Chat- ham Island. Fig. 3. Cellana strigilis flemingi Powell, Snares Islands. In figs. 2 and 3 the lateral, in semi-profile, and the median vestigial central only, are shown. 20 to 25 broadly rounded radial ribs, with an occasional weak interstitial one; surface smooth, apart from weak growth lines. Colour of exterior bluish-white, heavily blotched and streaked with light to dark-brown, the pattern frequently run- ning together, leaving elongated patches of the pale ground colour; interior pale amber, with the external pattern showing through in dark brown; spatula dark-brown in young shells, to clouded with buff or pale-brown in adults. Measurements (mm.)— length width height 57.5 45.00 20.00 paratype 41.0 29.00 10.50 holotype 35.0, 24.00 7.00 AWBP coll. 27.3 20.10 6.00 paratype 16.5 10.5 4.00 AWBP. coll. Synonymy— 1955 Cellana strigilis oliveri Powell, Dept. Sci. and Indust. Res., Cape Exped. Ser., Bull. no. 15, p. 73, pl. 5, figs. 48-50. Types—The holotype and paratypes are in the Dominion Museum, Wellington. Records—BOUNTY ISLANDS (Domin. Mus., Wellington); (AWBP coll.) Plate 169. Figs. 1-4. Cellana strigilis subspecies redimiculum (Reeve, 1854), New Zealand. Fig. 1. Stewart Island, 78 mm., AWBP coll. 40089. Figs. 2-4. Kartigi Beach, North Otago, 30-62. mm., AWBP. coll. 6874. Figs. 5-7. Cellana_ strigilis subspecies flemingi Powell, 1955, Snares Islands, southern islands of New Zealand, 30-50 mm., AWBP coll. 26611. Cellana strigilis subspecies redimiculum (Reeve, 1854) (Pl. 70, figs. 17-19; Pl. 169, figs. 1-4) Range—NEW ZEALAND: Stewart Island, eastern and western Otago, and east coast of South Island as far north as Kaikoura. Remarks—This is the common South Island mainland subspecies of strigilis, easily recognised by its orange-brown exterior, with pale blue ocellate flecks in the rib interstices, and inter- nally, by the dark brown radials, that do not anastomose at the margin. The centre of distri- bution is eastern Otago, where it is a common intertidal limpet. [01-742] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Cellana 189 Description—Shell rather large, up to 77 mm. (3 inches) in length, solid, broadly ovate, ele- vated, with the apex at about the anterior fourth or fifth. Sculpture consisting of about 20 strong rounded radial ribs, mostly with a much weaker radial in each interspace; the whole surface crossed by dense weak concentric growth lines. Colour of exterior orange-brown, with pale blue ocellate flecks and streaks in the radial inter- spaces; interior yellowish to greyish-brown, with a golden sheen, and the external rayed pattern showing through; spatula chestnut coloured, often clouded with a greyish callus. The margin bears a regular series of dark brown spots, mark- ing the terminal points of the external primary radials. Measurements (mm.)— (all A.W.B. Powell collection). length width height 77.0 65.0 33.0 Stewart Island 66.5 55.0 30.0 Kartigi OLED 43.0 21.0 Kartigi 48.0 36.0 14.0 Kartigi 30.0 22.0 10.0 Kartigi 35.5 27.5 10.5. Goose Bay Synonymy— 1854 Patella radians Gmelin, Reeve, Conch. Iconica, pl. 12, fivs. 25a, b. (non Gmelin, 1791). 1854 Patella redimiculum Reeve, Conch. Iconica, pl. 20, figs. 50a, b. 1873 Patella pottsi Hutton, Cat. Mar. Moll. N.Z., pp. 44. 1891 Helcioniscus redimiculum Reeve, Pilsbry, Man. Conch., vol. 13, p. 136, pl. 23, figs. 1,2,3,5. 1913 Helcioniscus strigilis (in part, non Hombron and Jac- quinot, 1841): Suter, Man. N. Z. Moll., p. 87. 1913 Helcioniscus redimiculum (in part): Suter, Man. N. Z. Moll., p. 85. 1927 Nacella redimiculum Reeve, Finlay, Trans. N. Z. Inst., vol. 57, pp. 337, 338. 1955 Cellana strigilis redimiculum Reeve, Powell Dept. Sci. and Indust. Res., Cape Exped. Ser., Bull. no. 15, p. 71. Types—The type of redimiculum is in the British Museum (Natural History). Records—NEW ZEALAND: South Island: Goose Bay, Kai- koura; Oamaru; Kartigi Beach, Otago; Waikouaiti, Otago; Portobello, Dunedin; St. Clair, Dunedin (all AWBP coll. 280); entrance to Milford Sound (Galathea Exped., Sta. 624); Henrietta Bay, Ruapuke Island, Foveaux Strait (Auck Inst.); Stewart Island (AWBP coll.); Blind Passage, Port Pegasus (Auck. Inst.). Cellana thomsoni Powell and Bartrum, 1929 (PI. 171, fig. 1) Range—New Zealand, lower Miocene. Remarks—The species is unlike any other New Zealand member of the genus, but bears some resemblance to the Japanese Recent toreuma Reeve. In the Japanese species, however, the radials are not so strongly or so numerously beaded. Description—Shell small, 14 mm. (9/16 of an inch) in length, but probably attained a much larger size, elongate-ovate, depressed, with the apex at about the anterior sixth. Sculpture con- sisting of about 36 primary narrowly rounded radials, and 1 to 3 secondary radials in the interspaces. The whole surface is crossed by very numerous concentric lamellose growth lines, that thicken to become knotted or beaded where they cross the radials. Measurements (mm.)— length width height 14.0 10.0 21.1 holotype Synonymy— 1929 Cellana thomsoni Powell and Bartrum, Trans. N. Z. Inst., vol. 59, p. 413, pl. 35, fig. 12. Types—The holotype, the only known specimen, is in the Geology Department, University of Auckland. Records—NEW ZEALAND: Waiheke Island, Oneroa Beds, Waitemata Group, Otaian Stage, lower Miocene. Plate 170. Juvenile colour patterns, by transmitted light, in New Zealand Cellana_ strigilis and subspecies. Fig. 1. C. strigilis redimiculum (Reeve), Oamaru, South Island. Fig. 2. C. strigilis flemingi Powell, Snares Islands. Fig. 3. C. strigilis strigilis (Hombron & Jacquinot), Auckland Islands. Fig. 4 C. strigilis oliveri Powell, Bounty Islands. Fig. 5. C.. strigilis bollonsi Powell, Antipodes Islands. The line is drawn through the apices [01-743] 190 Subgenus Cellana Plate 171. Fig. 1. Cellana thomsoni Powell & Bartrum, 1929, New Zealand, Waiheke Island, Auckland, Otaian, lower Miocene, 14 mm. holotype. Cellana cophina Powell, new species (PI. 172, fig. 1) Range—New Zealand. Cape Rodney, Hauraki Gulf (holotype), and coast % mile east of Goat Island, Cape Rodney (paratype). Motutapu Is- land, Auckland; all in either coarse sandstone or conglomerate, basal Waitemata Beds, Otaian, lower Miocene. Remarks—This strongly sculptured species is nearest allied to the Recent denticulata (Martyn, 1784), from which it differs in that both the radial ribs and the concentric cords are so strong that a coarse basket-weave effect results. Description—Shell rather large, 60 to 70 mm. (2%—2M inches) in length, narrowly ovate and of low profile, with the apex at about the anterior third. Sculpture very strong, like a coarse basket- weave, consisting of about 20 strong rounded radials, without intermediates, and crossed by closely spaced prominent cords that are much thickened where they cross the radials, but weak in the interspaces. Measurements (mm.)—Estimated size in paren- theses. length width height 59.0 (57.0) 43.5 (47.0) — (15.0) holotype 65.0 (70.0) 16.0 (16.0) paratype Types—Holotype and paratype in the collection of the New Zealand Geological Survey, Welling- ton. The type locality is Cape Rodney, Hauraki Gulf, New Zealand. A. W. B. Powell Patellidae Cellana taberna Powell, new species (Pl. 172, fig. 2) Range—New Zealand. Curiosity Shop, Rakaia, South Island, Waitakian greensands, lower Mio- cene. Remarks—This shell has distinctive sculpture, unlike that of any of the described species from New Zealand or elsewhere. The sculpture differs from that of C. cophina in that is a combination of large irregularly-oval, smooth blisters on the pri- mary radials with much weaker and more regular interstitial riblets that are cut into small, squarish nodes by deeply-incised concentric grooves. Description—Shell small, probably not adult, 23 mm. (% of an inch) in length, narrowly ovate, and of low profile, with the apex at the anterior third. Sculpture very strong, consisting of about 13 prominent rounded primary radials that de- velop large ovate smooth blisters, stronger pos- terior to the apex. Interstices with from 2 to 5 secondary radials, crossed by deeply incised con- centric grooves, cutting them into series of small rectangular nodes. Measurements (mm. )— length width height 22.5 17.25 6.25. holotype Type—The unique holotype is in the collection of the New Zealand Geological Survey, Welling- ton. The type locality is stated in the range above. Plate 172. Fig. 1. Cellana cophina new species. New Zealand, Cape Rodney, Hauraki Gulf, North Island, Otaian, lower Miocene; holotype, 55 (57) mim. Fig. 2. Cellana taberna new species. New Zealand, Curiosity Shop, Rakaia, South Island, Waitakian greensands, lower Miocene; holotype, 22.5 mi. [01-744] November 27, 1973 Genus Nacella Schumacher, 1817 Type Patella mytilina Helbling, 1779 This genus and its subgenus Patinigera are characteristic molluscs of Antarctic and Subant- arctic seas. They have their centre of distribu- tion in the Magellanic Province of southern South America from whence the seaweed-dwelling species in particular tend to spread eastward, being assisted to a considerable extent by the prevailing West Wind Drift that operates strongly in the Subantarctic Zone. Although Nacella and Patinigera are not members of the Indo-Pacific fauna, the recog- nised species are listed and briefly described here, since many of them do occur in waters to the south of both the Indian and Pacific Oceans, and at one location, Campbell Island, in the New Zealand faunal area, both Nacella (Patinigera) and the warmer-water derived Cellana flourish side by side. : Be = ba Plate 173. Figs. 1, 2. Nacella mytilina (Helbling, 1779), Mouth of Santa Cruz River, Patagonia, 27-29 mm., AWBP coll, 42389. Figs. 3, 4. Nacella kerguelenensis (E. A. Smith, ISv7). Fig. 3. Swain’s Bay, Kerguelen Island, 43 mm., AWBP coll, 30635. Fig. 4. Heard Island, 64 mm., AWBP coll. 40S61. INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Nacella 191 Both Nacella and Patinigera differ from all other Patellidae in the presence of an epipodial fringe, a scalloped lamellate flange that occupies a mid position between the edge of the foot and the gill cordon, except where it is interrupted by the head region. There is a link with Patella in that the gill cordon is complete, not interrupted by the head as it is in Cellana. The dentition, on the other hand, with its pair of centrals, alternating with a pair of laterals, is comparable with that of Cellana, not short, straight and bent back upon itself at the nascent end as it is in Patella. Thiele in 1929 proposed the subfamily Nacel- linae for Nacella, Patinigera and Cellana, but the epipodial fringe, characteristic of Nacella and its subgenus Patinigera, is not found in Cellana or in any other patellid genus. The radula, on the other hand, is very similar in all three of the above mentioned taxa, but very different from that of the Patellinae. Plate 174. Fig. 1. Kerguelen Nacella mytilina (Helbling), Island. Radula, from Thiele, in Troschel & Thiele, 1591, pl. 28, fig. 30. Fig. 2. Nacella (Patinigera) deaurata (Gmelin), Tuesday Bay. Radula, from Thiele, in Troschel & Thiele, 1591, pl. 28, fig. 32. Fig. 3. Nacella (Patinigera) terroris (Filhol), Campbell Island. Radula. [01-755] 192. Subgenus Nacella A. W. B. Powell Description—Shell rather small to moderately large, thin and fragile, typically elliptical, high- arched, with the apex strongly curved forward and downward, sometimes almost at the anterior end. The surface is smooth, or occasionally weakly radially ridged. Colour pale-olive to brownish, the apex coppery; inside silvery iri- descent to reddish bronze. The species live mostly attached to large seaweeds, and range from southern Chile and Argentina to the Ker- guelen Island. Synonymy— 1817 Nacella Schumacher, Essai d’un Noveau Systeme des Habitations, p. 179. Type, by subsequent designation, Gray, 1847: Patella mytilina Helbling, 1779. Nacella mytilina (Helbling, 1779) (Pl. 73, fig. 9; Pls. 173, 174) Range—Southern Chile, Straits of Magellan, Tierra del Fuego, Falkland Islands and Kerguelen Island. Description—Shell rather small, up to 43 mm. (1% inches) in length, elliptical, thin and fragile, with the apex almost at the anterior end. Usually the surface is almost smooth, but occasionally moderate radial ridges are developed, as. well as corrugations around the anterior margin. Colour greenish olive to light brownish, sometimes red- dish bronze at the apex, and the interior is silvery iridescent. Measurements (mm.)—(all A.W.B. Powell col- lection). length width height 43.0 26.0 18.0 Punta Arenas 34.5 23.0 11.0 Falkland Islands 27.0 18.0 7.5 Hermit Id., Cape Horn Synonymy— 1779 Patella mytilina Helbling, Abh. Privatges. Bohm., vol. 4, p. 104, pl. 1, figs. 5, 6. 1786 Patella mytiliformis Lightfoot, Cat. Portland Mus., p. 42. 1791 Patella conchacea Gmelin, Syst. Nat. ed. 13, p. 3708. 1817 Nacella mytiloides Schumacher, Essai Vers test., p. 179. 1819 Patella cymbularia Lamarck, Anim. sans Vert., vol. 6, p. 335. 1831 Patella cymbuloides Lesson, Voy. de la Coquille, p. 422. 1845 Patella hyalina Philippi, Arch. f. Naturg., vol. 11, p. 59. 1845 Patella cymbium Philippi, Arch. f. Naturg., vol. 11, p. 60. 1845 Patella vitrea Philippi, Arch. f. Naturg., vol. 11, p. 60. 1869 Nacella compressa Rochebrune & Mabille, Mission scient. Cap Horn, vol. 6, p. 98, pl. 5, fig. 9. 1913 Nacella falklandica’ Preston, Ann. Mag. Nat. Hist., ser. 8, vol. 11, p. 221, pl. 4, fig. 6. 1950 Nacella mytilina Helbling, Carcelles, Anales del Museo Nahuel Huapi, vol. 2, p. 52 (Kerguelen). 1951 Nacella| mytilina Helbling Powell, Discovery Rep., vol. 26, p. 80. Patellidae 1964 Nacella mytilina Helbling, Dell, Rec. Domin. Mus., vol. 4, no. 20, p. 273. Records—STRAITS OF MAGELLAN (type locality): Punta Arenas; St. Martin’s Cove, Hermite Island, Cape Horn. PAT- AGONIA: mouth of Santa Cruz River. FALKLAND ISLANDS (all AWBP coll.). KERGUELEN ISLAND: Swain’s Bay, inter- tidal and Antares Island, intertidal (BANZARE Sta. 48 and Sta. 61). Nacella kerguelenensis (E. A. Smith, 1877) (Pl. 73, fig. 10; Pl. 173, figs. 3, 4) Range—Kerguelen Island, Heard Island and Macquarie Island. Remarks—Dell (1964) has shown that the young stages of this species have the form of typical Nacella, and that in the adult the apex has moved back from near the front margin to a sub- central position. These adults, however, retain the light build of Nacella. It is possible that some of the lighter built Magellanic species of Pat- inella go through a Nacella stage also, but at present there is no evidence in support of this theory. The Macquarie Island record is based upon four beach shells, and none have been recorded since from that locality, so it is assumed that the original specimens may have drifted there upon floating kelp. Description—Shell thin and fragile, large, up to 80 mm. (3% inches) in length, broadly ovate, but decidedly narrowed anteriorly, rather eve- vated, and with the apex varying between near the front margin in juveniles to a subcentral position in adults. Sculpture consisting of weak radial folds. Colour dark purplish-brown, with the apex reddish-bronze; internally completely dark bronzy reddish-brown. Measurements (mm.)— length width height 80.0 65.5 33.0 Heard Island; Dell, 1964 67.0 57.0 24.0 Royal Sound, Kerguelen 46.0 37.0 15.6 Royal Sound, Kerguelen Synonymy— 1877 Patella (Patinella) kerguelenensis E. A. Smith, Phil. Trans Roy. Soc., London, vol. 168, p. 177, pl. 19, figs. 13, 13a. (Kerguelen Island). 1886 Patella kerguelenensis Smith, Watson, Challenger Rep., vol. 15, p. 27. 1908 Patinella kerguelenensis Smith, Strebel, Schwed. Sud- pol. Exped., Zool., vol. 6, p. $3. 1916 Nacella kerguelenensis Smith, Hedley, Aust. Ant. Exped. 1911-1914, ser. C, vol. 4, pt. 1, p. 44. Macquarie Island. 1957 Patinigera kerguelenensis Smith, Powell, B.A.N.Z. Ant. Res. Exped., vol. 6, p. 126. 1964 Nacella kerguelenensis Smith, Dell, Rec. Domin. Mus., vol. 4, no. 20, p. 276. [01-756] November 27, 1973 Subgenus Patinigera Dall, 1905 Type: Patella magellanica Gmelin, 1791 Shells of the subgenus are mostly more solid than those of typical Nacella. They have the apex well back from the anterior end, sometimes being subcentral in position. The interior is always with a bronze lustre. As in typical Nacella, the gill cordon is complete and there is a_well- developed epipodial fringe. The radula shows no important differences. The subgenus is more widely distributed than is typical Nacella. It extends up the western coast of South America as far as Valparaiso, and southward to the subantarctic islands and Ant- arctica. It also occurs at Macquarie Island, and reaches its furthest north location in the New Zealand faunal region at Campbell Island, D2. 30 Oo. Synonymy— 1871 Patinella Dall, Proceedings of the Boston Society of Natural History, vol. 14, p. 53. Type by original des- ignation: Patella magellanica Gmelin, 1791. 1905 Patinigera Dall, Nautilus, vol. 18, no. 10, nom. nov. pro Patinella Dall, 1871, non Gray, 1545. Nacella clypeater (Lesson, 1831) (PI. 73, fig. 13; Pl. 175, figs. 1,2) Range—Chile, to as far north as Valparaiso. Remarks—The species is easily recognised by its nearly circular outline. Description—Shell of moderate size, up to 61 mm. (2% inches) in length, rather depressed and almost circular in outline, with the apex sub- central. Sculpture consisting of very numerous, regular, narrow, low rounded, radial ribs. Colour dull reddish-brown, the ribs paler; interior silvery to pale bronze, with the spatula area irregularly blotched with dark reddish-brown. Measurements (mm.)— length width height 61.0 58.0 14.5 Chile; Pilsbry, 1891 56.5 51.0 17.0 Chile 47.0 43.5 14.5 Chile 37.0 32.0 9.0 Valparaiso INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Nacella 193 Synonymy— 1831 Patella clypeater Lesson, Voy. Coquille, Zool., vol. 2, p. 419. 1854 Patella clypeater Lesson, Reeve, Conch. Iconica, pl. 16, figs. 38 a, b. “Monterey, California,” in error. 1891 Nacella (Patinella) clypeater Lesson, Pilsbry, Man. Conch., vol. 13, p. 122, pl. 50, figs. 40-43. Nacella concinna (Strebel, 1908) (Pl. 176, figs. 1-5) Range—South Georgia, South Orkneys, South Shetlands, Bouvet Island, Seymour _ Island, Paulet Island, Wandel Island, Anvers Island and Petermann Island. Remarks—Unfortunately the well-known name of this species, Patella polaris Hombron & Jac- quinot, 1841, falls as a homonym of the same combination of Roéding, 1798. However there is a substitute name available, in Patinella polaris concinna Strebel, 1908, from South Georgia, and this name, concinna, may be used specifically, since there appears to be no real difference be- tween the shallow-water ‘polaris’ and the deeper- water concinna, other than a gradual tendency towards lower profile, lighter build, more clear- Plate 175. Figs. 1, 2. Nacella (Patinigera) clypeater (Lesson, 1831), Chile, 44-53 mm., AWBP coll. 46145. Figs. 3, 4. Na- cella (Patinigera) deaurata (Gmelin, 1791), Falkland Islands. 60 mm., AWBP coll. 632. [01-761] 194. Subgenus Patinigera A. W. B. Powell Patellidae cut ribbing and paler coloration as the depth increases. Strebel’s concinna, described as a Patinella, does not conflict with the Japanese acmaeid that was originally described as Patella concinna Lischke. The species lives from the intertidal zone down to 110 metres. Description (shallow-water ‘polaris’ form)— Shell moderately large, up to 60 mm. (2% inches) in length, elongate ovate, rather thin, moderately elevated, with the apex between central and the anterior third. Sculptured with distant weak radial ribs in young shells, but the ribbing be- comes subobsolete to obsolete in the adult. Col- our, externally pale brownish; internally very dark bronzy-brown, almost black, the spatula sometimes a paler chestnut-brown. Description (deeper-water typical form)—Shell usually small, 20 to 32 mm. (% to 14 inches) in length, thin and fragile, elongate ovate, moder- ately elevated and with the apex at about the anterior third. Sculptured with about 28 to 30 narrow radial .ribs, crossed by dense fine lamel- lose growth lines. Colour buff, sparingly blotched in reddish brown; interior cream, shining, vary- ingly maculated with pale reddish brown. Some larger examples, approaching the larger shallow- water form in size, tend to flatten out at the margin, towards which the radial sculpture be- comes subobsolete. Plate 176. Figs. 1-5. Nacella (Patinigera) concinna (Strebel, 1908). Figs. 1, 2. (coneinna form), East Cumberland Bay, South Georgia, 27 metres, 27 mm., AWBP coll. Fig. 3. East Cumberland Bay, 24-30 metres, 41 mm., AWBP coll. 26831. Figs. 4, 5. (polaris form), Melchior Island, Schollaert Channel, Palmer Archipelago, 4-10 metres, 58 mm., AWBP coll. 52493. Figs. 6, 7. Nacella (Patinigera) delesserti (Philippi, 1549), Marion Island, south western Indian Ocean, 35-53) mm., AW BP coll. 52491. [01-762] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Nacella 195 Measurements (mm.)— length width height 58.0 42.0 19.0 Palmer Archipelago, 4-10 metres: (‘polaris’ form) 44.0 31.0 18.5 S. Orkneys (‘polaris’ form) 42.0 29.0 9.0 S. Georgia, 27 metres: (intermediate form) 31.0 21.5 11.0 S. Georgia (‘polaris’ form) 29.0 20.0 8.0 S. Georgia, 18 metres: (typical concinna) 25.0 17.0 6.25 S. Georgia (typical concinna) Synonymy— 1841 Patella polaris Hombon & Jacquinot, Ann. Sci. Nat. Zool., vol. 16, p. 191 (non Réding, 1795). 1886 Patella polaris H. and J. Martens and Pfeffer, Moll. Siid-Georgien, J. hamb. wiss. Anst., vol. 3, p. 101, pl. 2, figs. 11-13. 1891 Nacella (Patinella) polaris H. and J. Pilsbry, Man. Conch., vol. 13, p. 120, pl. 49, figs. 21-27. 1908 Patinella polaris H. and J. Strebel, Wiss. Ergeb. schwed. Siidpolar-Exped. (1901-3), vol. 6, p. 81, pl. 5, fig. 77. 1908 Patinella polaris concinna Strobel, Wis. Ergeb. schwed. Stidpolar-Exped. (1901-3), vol. 6, p. $2, pl. 5, figs. 76 a-e, 78 a, b. 1951 Patinigera polaris H. and J. Powell, Discovery Rep., vol. 26, p. 82. 1951 Patinigera polaris concinna Strebel, Powell, Discovery Rep., vol. 26, p. 83. Records—SOUTH GEORGIA (type of ‘polaris’): Cumber- land Bay, 15-25 metres (type of concinna); East Cumberland Bay, 18-110 metres; Moltke Harbour, in rock pool; Strom- ness Harbour, 26-35 metres; Undine Harbour, 18-27 metres. SOUTH ORKNEYS: Signy Island, 18-27 metres; Normanna Strait, 24-36 metres (‘Discovery II’; Powell, 1951). SOUTH SHETLANDS: Deception Island, 5-60 metres; Nelson Island, shore; Livingston Island, shore; Wilhelmina Bay, Danco Land, 1-8 fathoms (‘Discovery I’; Powell, 1951). PALMER ARCH- IPELAGO: Melchior Island, 4-10 metres. Bouvet Island, 40-45 metres (‘Discovery I’; Powell, 1951). Seymour, Paulet, Wandel, Anvers and Petermann Islands (Strebel, 1908; Lamy, LOU): Nacella deaurata subspecies deaurata (Gmelin, 1791) (Pl. 73, fig. 11; Pls. 174, 175) Range—Southern Patagonia, Straits of Ma- gellan, Tierra del Fuego and Falkland Islands. Remarks—The_ species is nearest allied to magellanica which is more broadly ovate in out- line and lacks nodulation of the radials. Description—Shell of moderate size, up to 61 mm. (2% inches) in length, rather solid, tall conical, narrowly ovate, and with the apex at about the anterior third. Sculpture consisting of from 36 to 40 strong radial ribs, which are ren- dered strongly scabrous to nodular by numerous overriding concentric lamellose lirae. Colour yellowish-brown to reddish-brown, tending dark reddish-brown to bronze over the apical area. Interior silvery with a pinkish lustre, more or less rayed and mottled with reddish-bronze, the spatula and spotted marginal border dark red- dish-brown. Measurements (mm.)— length width height 61.0 43.0 27.0 Falkland Islands 57.0 43.0 24.0 Falkland Islands 48.0 34.0 19.5. Falkland Islands Synonymy— 1784 Patella aenea Martyn, Univ. Conch., vol. 1, fig. 17 (invalid). 1791 Patella deaurata Gmelin, Syst. Nat. ed. 13, p. 3719, based upon Martini-Chemnitz, Conch. Cab., vol. 10, p. 327, pl. 168, figs. 1616 a, b. 1854 Patella varicosa Reeve, Conch. Iconica, pl. 11, figs. 21 a-c. 1885 Nacella — strigatella| Rochebrune and = Mabille, Bull. Soc. Phil. Paris, ser. 7, vol. 9, p. 110. 1891 Nacella (Patinella) aenea Martyn, Pilsbry, Man. Conch. vol. 13, p. 115, pl. 46, figs. 28-36. 1913 Helcioniscus bennetti Preston, Ann. Mag. Nat. Hist. ser. 8, vol. 11, p. 221, pl. 4, fig. 7. 1951 Patinigera aenea Martyn, Powell, Discovery Rep. vol. 26, p. 82. Nacella deaurata form delicatissima (Strebel, 1907) (Pl. 178, figs. 3, 4) Range—Straits of Islands. Remarks—This is a small thin shell of low pro- file with delicately squamose ribs, and of pale colour with a few rays and streaks of reddish- brown at most. The writer has insufficient ma- terial to properly evaluate this shell which may prove to grade into the typical species. It occurs from 5 to 50 fathoms. Measurements (mm.)— Magellan and Falkland length width height 46.7 36.9 15.8 Strebel, 1908, pl. 5, fig. 75 21.4 16.6 5.7 Strebel, 1905, p. 145. 15.0 10.75 3.25. Eddystone Rock, Falklands, 115 metres Synonymy— 1907 Patinella delicatissima Strebel, Zool. Jahrb. Abt. Syst., Jena, vol. 25, p. 145, pl. 5, figs. 71, 72, 74, 75. 1908 Patinella delicatissima Strebel, Wiss. Ergeb. schwed. Siidpolar-Exped., vol. 6, pt. 1, pl. 1, figs. 75, 75a. 1951 Patinigera delicatissima Strebel, Powell, Discovery Rep., vol. 26, p. 82. Records—STRAITS OF MAGELLAN: 20-30 fathoms (type locality); Uschuaia, Tierra del Fuego, 1-2 fathoms (Strebel, 1908). FALKLAND ISLANDS: off Eddystone Rock, East Falkland Islands, 115 metres; entrance to Port Stanley, 10-16 metres; Sparrow Cove, Port William, 10.5-16 metres (Dis- covery 11, Sta. 51, 52 and 56). [01-763] 196 Subgenus Patinigera A. W. B. Powell Patellidae Nacella delesserti (Philippi, 1849) (Pl. 176, figs. 6, 7) Range—Marion Island (between South Africa and Antarctica). Remarks—The writer has only two examples of this species available but they seem to represent a distinct species, characterised by an elongate ovate outline and simplicity of both sculpture and colour pattern. According to Hedley (1916, Aust. Ant. Exped., ser. C, vol. 4, pp. 42, 43) delesserti was based upon an immature shell of only 22 mm. in length. Reeve (1854, Conch. Iconica, sp. 40, pl. 17) named a shell from unknown locality, Patella ferruginea, basing it apparently, upon a manuscript species of Sowerby, and with Patella delesserti Philippi cited in its synonymy. How- ever Reeve’s figures are unlike any shell I have seen and certainly bear no resemblance to the Marion Island limpet. Description—Shell of moderate size, up to 54 mm. (2% inches) in length, elongate ovate, more narrowly rounded anteriorly and of moderate elevation, with the apex between the anterior Plate 177. Figs. 1, 2. Nacella (Patinigera) fuegiensis (Reeve, 1855), “Tierra del Fuego. Falkland Islands;;, from Reeve, 1855, pl. 28, fig. 73. Figs. 3, 4. Nacella (Patinigera) edgari (Powell, 1957), Royal Sound, Kerguelen Island, in fish trap, 34 mm., AWBP coll. third and fourth. Sculpture consisting of about 24 low, carinated to rounded, radial folds and an occasional intermediate, the whole — surface densely crossed by weak lamellose growth lines. Colour of exterior greyish-white, with most of the primary radials dark reddish-brown. Interior heavily blotched and radially streaked in dark reddish-bronze. Spatula very large. Measurements (mm.)— length width height 53.0 40.0 16.0 Marion Island 38.0 27.5 13.0 = Marion Island Synonymy— 1849 Patella delesserti Philippi, Abbild. Conch. vol. 3, pt. 4, p. 9, pl. 1, fig. 5. Nacella edgari (Powell, 1957) (Pl. 177, figs. 3, 4; pl. 179) Range—Kerguelen Island. Remarks—The adult shell is very thin and frag- ile, almost flat to slightly concave, with an animal too large to allow the shell tight contact with the surface of the kelp. Young examples sometimes occur on rocks, and such have a slight elevation, as well as crisp narrow radials, but these become Plate 178. Figs. 1, 2. Nacella (Patinigera) magellanica (Gmelin, 1791), Straits of Magellan, 51 mm., AWBP. coll. 45751). Figs. 3, 4. Nacella (Patinigera) deaurata subspecies delicatissima (Strebel, 1907), Falkland Islands, 115 metres, 16 mm., AWBP coll. [01-764] November 27, 1973 subobsolete as the shell grows and flattens out at the edges. It is associated with kelp from inter- tidal to about 55 metres. Description—Shell elongate-ovate, of moderate size, up to 51 mm. (2 inches) in length, very thin and fragile, and very depressed, only the apical area slightly raised, and much of the remaining dorsal surface actually concave. Sculpture con- sisting of many radial folds, that are narrow at first but soon become broad and low, almost disappearing towards the margin in adults. The whole surface crowded with sharp concentric lamellae that undulate as they cross the radials. Colour dull-slate externally, tinged with bronze at the apex; internally, iridescent bluish grey, with the spatula and an irregular marginal border diffused reddish brown. Measurements (mm.)—(All station numbers are of the British, Australian and New Zealand Antarctic Research Expedition, 1929-1931). length width height 51.0 33.0 8.0 holotype; Royal Sound, 20 metres. Sta. 5 43.0 30.5 3.5. Colbeck Passage, 20 metres. Sta. 55a. 40.0 27.5 4.5 Royal Sound, 1-5 metres. Sta. Sob. 38.0 26.5 10.5. Swain’s Bay, intertidal. Sta. 48. Synonymy— 1877 Patella (Patinella) fuegiensis Reeve, E. A. Smith, Phil. Trans. Roy. Soc. London for 1879 (issued separately 1877), vol. 168, p. 180, pl. 9, figs. 14, 14a. Not Reeve, 1855. 1891 Nacella (Patinella) fuegiensis Reeve, Pilsbry, Man. Conch. vol. 13, p. 121 (in part, pl. 49, figs. 28, 29 only). Not Reeve, 1855. 1957 Patinigera fuegiensis edgari Powell, B.A.N.Z. Ant. Res Exped., vol. 6, pt. 7, p. 127, text figs. 1, la, Ib. Types—The holotype and paratypes are at pres- ent in the Auckland Museum. Records—KERGUELEN ISLAND, thirteen B.A.N.Z.A.R-E. Stations from in and around Royal Sound, ranging from low tide to 55 metres. The type locality is Port Jeanne d’Arc, Royal Sound, 20 metres, along the lower edge of the kelp belt. ee Plate 179. Nacella (Patinigera) edgari (Powell, 1957), Grotto Bay, Port Jeanne d’ Arc, Kerguelen Island, 10 metres, 37.5 x 25.5 x 4mm. In profile to show the very slight elevation of the shell. INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Nacella 197 Nacella flammea (Gmelin, 1791) (PI. 181) Range—Straits of Magellan. Remarks—This shell resembles fuegiensis in its narrowly oval outline, moderate elevation, and light build, but differs in sculpture, the radials being almost obsolete, represented at most by broad very weak radial folds towards the margin. The colour pattern is of broad irregular axial streaks upon a whitish ground. Description—Shell elongate-ovate, of light build, and of moderate size, up to 40 mm. (1% inches) in length, moderately elevated, and with the apex at between the anterior third and fourth. Surface relatively smooth, just a few, almost obsolete, broad low radial folds over the posterior half of the shell, and only towards the outer mar- gin. The only other sculpture consists of very faint concentric growth lines. External colour pattern of broad, flexuous, dark reddish brown axial streaks upon a whitish ground; internally, the colour pattern is the same, except for a bright chestnut spatula. Measurements (mm. )— length width height 39.0 27.8 10.0 Strebel, 1907, p. 145 33.) 24.0 9.5 St. of Magellan 27.0 18.7 7.0 St. of Magellan Synonymy— 1791 Patella flammea Gmelin, Syst. Nat., ed. 13, p. 3716; based upon Martini-Chemnitz, Conch. Cab., 1, pl. 5, fig. 42. 1907 Patinella flammea: Strebel, Zool. Jahrb., 25 (1), p. 145, pl. 5, fig. 73. Nacella fuegiensis (Reeve, 1855) (Pl. 177, figs. 1, 2) Range—Tierra del Fuego, Falkland Islands, Petermann Island and South Georgia. Remarks—This is a thin-shelled, elongate-ovate species, with numerous, weak, almost smooth, radial ribs. It is of pale greenish ground colour, partially to almost entirely blotched with bronzy reddish brown. The somewhat similar deaurata is of stouter build and has the radials strongly scabrous. Description—Shell elongate-ovate, of moderate size, up to 50 mm. (2 inches) in length, thin and fragile, moderately elevated and slightly laterally compressed. Sculpture consisting of numerous, narrow, sharply raised radials, in early stages of growth, but these tend to become broader and [01-765] 198 Subgenus Patinigera A. W. B. Powell Patellidae lower towards maturity; the whole crossed by very dense sharp concentric lirae. Colour green- ish grey, more or less blotched with reddish brown, and the apical area with a bronze lustre. Interior iridescent bronzy-brown, rayed_ with darker brown, corresponding to the external sculptural interspaces. Synonymy— 1855 Patella fuegiensis Reeve, Conch. Iconica, pl. 28, figs. 73 a, b. 1891 Nacella (Patinella) fuegiensis Reeve, Pilsbry, Man. Conch., vol. 13, p. 121, pl. 49. figs. 30, 31 (non figs. 28, 29). Nacella magellanica subspecies magellanica (Gmelin, 1791) (PI. 73, figs. 14, 15; Pl. 178, figs. 1, 2) Range—Tierra del Fuego, Straits of Magellan, Patagonia and Falkland Islands. Remarks—This is the common limpet of the Magellanic Region, and it is easily recognised by its roundly oval shape, high-conical profile, nearly central apex, and = strong unsculptured radial ribbing. Description—Shell of moderate size, up to 65.6 mm. (2% inches) in length, rather solid, roundly ovate and high-conical, with the apex erect and near central, strongly and regularly sculptured with relatively few bold rounded radials that deeply corrugate the margin. The concentric sculpture is confined to weak growth lines that do not render the radials either scabrous or beaded. Colour of exterior variable, pale reddish brown to greenish grey or brown, occasionally with broad dark-brown radial bands. Interior metallic- brown or leaden with the spatula bronzy-chest- nut, and either a continuous or spotted marginal border of very dark-brown, the brown spots, when present, corresponding to the external ribs. Measurements (mm.)— length width height 65.6 58.3 41.5 Strebel, 1907, fig. 94a 53.5 45.5 31.0 Straits of Magellan 45.0 37.5 21.5 Punta Arenas 33.0 27.0 18.0 Straits of Magellan 26.0 21.0 13.5. Straits of Magellan Synonymy— 1791 Patella magellanica Gmelin, Syst. Nat. ed. 13, p. 3703, based upon Gault. pl. 9, fig. E, and Martini-Chem- nitz, Conch. Cab. vol. 1, pl. 5, figs. 40 a, b. 1854 Patella) magellanica Gmelin, Reeve, Conch. Iconica, pl. 10, figs. 19 a, b. 1854 Patella atramentosa Reeve, Conch. Iconica, pl. 17, figs. 4] a, b. 1885 Patella) meridionalis Rochebrune and = Mabille, Bull. Soc. Phil., Paris, ser. 7, vol. 9, p. 109. 1885 Patella metallica Rochebrune and Mabille, Bull. Soc. Phil., Paris, ser. 7, vol. 9, p. 109. 1885 Patella pupillata Rochebrune and Mabille, Bull. Soc. Phil., Paris, ser. 7, vol. 9, p. 110. 1885 Patella tincta} Rochebrune and Mabille, Bull. Soc. Phil., Paris, ser. 7, vol. 9, p. 110. 1891 Patinella aenea var. magellanica Gmelin, Pilsbry, Man. Conch., vol. 13, p. 119, pl. 44, figs. 9-17; pl. 43, figs. 1-6. 1907 Patinella| magellanica-atramentosa Strebel, Zool. Jahrb. Abt. Syst. Jena, vol. 25, p. 146, pl. 6, figs. 86-88; pl. 7, figs. 91, 92, 94, 95. 1907 Patinella aenea var. minor Strebel, Zool. Jahrb. Abt. Syst. Jena, vol. 25, p. 137, pl. 5, figs. 67 a-d. 1951 Patinigera magellanica Gmelin, Powell, Rep., vol. 26, p. 81. Discovery Nacella magellanica subspecies venosa (Reeve, 1854) (PI. 180, figs. 1-4) Range—Chiloe Island, Chile. Remarks—Both venosa and chiloensis, from the same locality, Chiloe Island, appear to represent but one form, a roundly ovate, high-conical, thin- shelled, sub-obsoletely sculptured variant of magellanica, which, as suggested by Dell (l.c., 1964) may be a regional subspecies. Unfortun- ately the writer has no material available upon which to make further comment. Plate 180. Nacella venosa (Reeve, 1854) Chiloe Island, Chile. Figs. 1, 2. Patella venosa Reeve, 1854, Conch. Iconica, pl. 10, figs. 1S a, b. Figs. 3, 4. Patella chiloensis Reeve, 1855, Conch. Iconica, pl. 33, figs. 98 a, b. (Patinigera) magellanica subspecies [01-766] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Nacella 199 Description—(original, for venosa) “Shell ovate, convex, rather high with age, a little contracted in front, in the young shell radiately ribbed, ribs small, rather distant, more or less obsolete with age, varicose near the margin; transparent-white, irregularly veined with chestnut-purple, veins bifurcated at the margin, deeply stained with purple-chestnut in the interior.” Reeve’s chiloen- sis is essentially similar in shape, height, sculp- ture and position of the apex, but differs in color- ation, a minor point, in being irregularly stained and streaked in smoky-black. Synonymy— 1854 Patella venosa Reeve, Conch. Iconica, pl. 10, figs. 18 a-c. 1855 Patella chiloensis Reeve, Conch. Iconica, pl. 33, figs. 98 a, b. 1964 Patinigera magellanica venosa Reeve, Dell, Rec. Domin. Mus., vol. 4, no. 20, p. 273. Nacella macquariensis Finlay, 1927 (Pl. 182, figs. 1, 2) Range—Macquarie and Heard Islands. Remarks—Compared_ with terroris, macquar- iensis varies greatly both in outline and in height; also it has fewer and stronger primary radials, a tendency to be longer and narrower, and the spat- ula is always clearly defined, often heavily callused. In terroris the shape is constantly broadly ovate and the spatula is never clearly defined. Description—Shell moderately large, up to 63 mm. (2% inches) in length, strong but of light build, mostly narrowly ovate and elevated, but varying to rather broadly ovate and depressed. Sculpture consisting of from 32 to 35 broadly rounded primary radial ribs, plus narrower secondary intermediate radials, mostly over the posterior half of the shell; the whole surface crowded with low concentric growth lamellae, that do not thicken to any extent on the crests of the radials. Colour, externally, olive to chest- nut-brown; internally, diffused and __ strongly rayed in bronzy reddish brown, the spatula well- marked, variously blotched with reddish brown, and often almost completely white-callused. Measurements (mm. )— length width height 63.0 51.9 31.2. Garden Cove; Dell, 1964 58.0 46.0 22.0 Macquarie Island 43.0 33.0 14.0 Hurd Point 41.0 35.0 17.0 Hurd Point 41.0 30.0 12.5 Hurd Point Synonymy— 1913 Nacella fuegiensis Reeve, Suter (in part), Man. N. Z. Moll., p. 77. Not Reeve, 1855. 1916 Nacella delesserti Philippi, Hedley, Aust. Ant. Exped., ser. C, vol. 4, pt. 1, p. 42, pl. 6, figs. 65-69. Not Phil- ippi, 1849. 1927 Nacella macquariensis Finlay, Trans. N. Z. Inst., vol. 57, p. 337. 1955 Patinigera macquariensis Finlay, Dell, Rec. Domin. Mus., vol. 4, no. 20, p. 274. Types—The type series, based upon Hedley’s 1916 figures (pl. 6, figs. 65-69), is in the Aus- tralian Museum, Sydney. Records—MACQUARIE ISLAND (AWBP. coll.); Hurd Point (AWBP. coll.); Aerial Cove and Garden Cove (Dell, 1964). HEARD ISLAND: Atlas Cove and Cape Gazert (Dell, 1964). Nacella terroris (Filhol, 1880) (Pl. 73, fig. 12; Pls. 174, 182) Range—Campbell Island, only, New Zealand subantarctic. Remarks—This species marks the furthest north attained by this cold water genus in the New Zealand area, the latitude of Campbell Island being 52° 33’ S. It is also interesting that at this same location, Cellana strigilis is abundant, in turn marking the most southerly occurrence of that warm water Indo-Pacific genus. Description—Shell moderately large, up to 57 mm. (24 inches) in length, strong but of light build, rather broadly ovate, and_high-conical, with the apex at about the anterior third. Sculp- ture consisting of about 50 narrow radial ribs, about 40 of them primary, the remaining ones being short and interpolated around the posterior margin. The whole surface is crossed by dense, crisp, undulating concentric growth lamellae, Plate 181. Nacella (Patinigera) flammea (Gmelin, 1791), Straits of Magellan, 27 mm. & 33.5 mm., AWBP coll. 46064. [01-767] 200 Subgenus Patinigera that thicken where they cross the radials. Colour of exterior reddish to greenish-brown, without markings; interior, pale purplish grey, densely radiately lined in reddish-purple, and with irreg- ular blotches of bronzy reddish brown over the central area, which lacks a clearly defined spatula. Animal—As in other members of the genus, there is a prominent scalloped epipodial fringe that lies between the edge of the foot and the branchial cordon. This epipodial fringe is inter- rupted by the head but the branchial cordon is not. Radula—Formula 3 + 1 +(1+0+1)+:> 1 + 3. Mod- erately long and loosely coiled, in several volu- tions, on the right hand side of the animal, when viewed from above. This feature recalls the radula of Cellana, except that in that genus it is still longer and has more coils. Measurements (mm.)—(all A. W. B. Powell, coll.). length width height 50.0 39.0 24.0 Perseverance Harbour 57.0 46.5 27.0 Perseverance Harbour 46.0 36.0 15.0 Perseverance Harbour 39.0 31.5 17.0 Perseverance Harbour 33.25 26.75 15.0 Perseverance Harbour Synonymy— 1880 Patella terroris Filhol, Compt. Rend., vol. 91, p. 1095. 1885 Patella terroris: Filhol, Mission !He Campbell, p. 529. 1913 Nacella (Patinigera) illuminata (non Gould): Suter (in part), Man. N. Z. Moll., p. 77. 1955 Patinigera terroris: Powell, D.S.I.R. Cape Exped. Ser., Bull. 15, p. 69. Types—The type specimens are in the Muséum National d’Histoire Naturelle, Paris. A. W. B. Powell Patellidae Records—CAMPBELL ISLAND (type), New Zealand sub- antarctic: Perseverance Harbour, on rocks at low tide (Auck. Mus.; AWBP coll.). Plate 182. Figs. 1, 2. Nacella (Patinigera) macquariensis Finlay, 1927, Hurd Point, Macquarie Island, 40 mm., AWBP coll. 42864. Figs. 3, 4. Nacella (Patinigera) terroris (Filhol, 1880), Perseverance Harbour, Campbell Island, 52> mm., AWBP coll. 26164. [01-768] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 INDEX TO PATELLIDAE NAMES IN VOL. 3, NO. 15 Looseleaf subscribers should keep this index at the beginning of the family Patellidae. The family begins on page 75 (loose- leaf p. 01-551). In this index, the number following the name refers to the pagination found at the top of the page in vol. 3, no. 15. The column at the right is the looseleaf pagination. All new names proposed in this number are in boldface type. Acmaeidae characters, 76 aculeata Reeve, 137 adansonii Dunker, 111 adspersa Bucquoy, Dautzenberg & Dollfus, 99 aenea Martyn, 195 affinis Reeve, 163 alba Christiaens, 98 alba Venison Woods, 135 albanyana, Turton, 126 alboguttata Turton, 119 alboradiata Turton, 116 amphitrite Turton, 125 amuritica Wilckens, $2 amussitata Reeve, 163 analogia Iredale, 173 Ancistromesus Dall, 141 angulosa: Turton, 126 Ansates Sowerby, 144 antipodum Smith, 176 antipodum E. A. Smith, 182 apicina Lamarck, 110 approximata, Turton, 145 ardosiaea Hombron & Jacquinot, 167 argentata Sowerby, 166 argentea Quoy & Gaimard, 182 argenvillei Krauss, 123 argyropsis Lesson, 182 ariel Iredale, 170 arrecta Iredale, 130 [looseleaf ] 01-552 01-66] 01-615 01-583 01-763 Ory. 01-582 01-662 01-652 01-637 01-630 01-651 01-558 Ol-717 01-727 01-675 01-652 (1-682 (1-730 01-736 01-614 (1-687 01-721 01-720 01-736 01-649 01-736 01-724 01-654 articulata Reeve, 151 aspera Lamarck, 52 aspera Réding, 82 assimilans Turton, 116 aster Reeve, 149 astrolepas Réding, 111 athletica Bean, 82 atramentosa Reeve, 198 aurea Martel, 95 aurorae Fleming, 133 azorica Nuttall, 100 badia Gmelin, 111 barbara Linnaeus, 124 barbata Lamarck, 125 baudonii Drouet, 96 becki Turton, 119 bennetti Preston, 195 bimaculata Montagu, 144 bollonsi Powell, 185 bombayana E. A. Smith, 149 boninensis Pilsbry, 16] bonnardi Payraudeau, 99 brunescens Turton, 125 caerulea Linnaeus, 99 caerulea Quoy & Gaimard, 113 candei d’Orbigny, 103 canescens Gmelin, 112 capensis Gmelin, 149 carpentariana Skwarko, 172 Cellana A. Adams, 147 [01-501] Index 201 [looseleaf ] 01-705 01-558 01-558 01-630 01-703 01-615 01-558 01-766 01-579 01-657 01-554 01-615 01-650 01-651 01-580 01-637 01-763 01-682 01-739 01-703 01-715 01-583 01-651 01-583 01-617 01-597 01-616 01-703 01-726 01-701 202. Index cernica H. Adams, 155 ceylanica E. A. Smith, 149 chapmani Tenison Woods, 137 chathamensis Pilsbry, 186 chiloensis Reeve, 199 chitonoides Reeve, 128 citrullus Gould, 103 clathratula Reeve, 168 clypeater Lesson, 193 cochlear Born, 121 coeruleata da Costa, 144 cognata Bucquoy, Dautzenberg & Dollfus, 99 communis Brown, 95 compressa Linnaeus, 117 compressa Rochebrune & Mabille, 192 conchacea Gmelin, 192 conciliata Iredale, 165 concinna Strebel, 193 concolor Krauss, 103 conica Anton, 95 conica Brown, 95 conspicua Philippi, 113 conspicua Turton (Helcion), 145 conspicua Turton (Patella), 106 constellata Sowerby, 106 conus Réding, 95 cooperi Powell, 139 cophina Powell, n. sp., 190 cornea Potiez & Michaud, 144 cornea Helbling, 144 corrugata Oliver, 176 Costatopatella Pallary, 95 costoso-plicata Reeve, 96 craticulatus Suter, 174 crenata Gmelin, 99 cretacea Reeve, 130 cudmorei Chapman & Gabriel, 172 cuprea Reeve, 166 cymbium Philippi, 192 Cymbula H. and A. Adams, 117 cymbularia Lamarck, 192 cymbuloides Lesson, 192 cypria Gmelin, 125 cypridium Perry, 144 deaurata Gmelin, 195 decemcostata E. A. Smith, 126 decora Philippi, 182 decorata Turton, 119 deformis K. Martin, 153 A. W. B. Powell Patellidae [looseleaf] 01-709 01-703 01-661 01-740 01-767 01-654 01-597 01-722 01-761 01-643 01-682 01-583 01-579 01-635 01-756 01-756 01-722 01-761 01-597 01-579 01-579 01-617 01-687 01-600 (1-600 01-579 01-663 01-744 (01-682 (1-682 01-730 01-579 01-580 01-728 01-583 01-654 01-726 01-720 01-756 01-635 01-756 01-756 01-651 01-682 01-763 (1-652 01-736 01-637 (1-707 [looseleaf] delesserti Philippi, 196. 01-764 delicatissima Strebel, 195 01-763 denseplicata Turton, 119 01-637 densestriata Turton, 119 01-637 denticulata Martyn, 177 01-731 depressa Pennant, 82 01-558 depsta Reeve, 106 (1-600 dimenensis Philippi, 137 01-661 dirus: Iredale, 176 01-730 discrepans Pilsbry, 155 01-712 divergens Pilsbry, 163 01-717 dunkeri Krauss, 145 01-687 earlii Reeve, 182 01-736 echinulata Krauss, 116 01-630 edgari Powell, 196 01-764 electrina Reeve, 83 01-559 elliptica Fleming, 144 01-682 elevata Jeffreys, 95 01-579 elongata Fleming, 144 01-682 enneagona Reeve, 150 01-704 eucosmia Pilsbry, 147 01-701 eudora Iredale, 153 01-707 exarata Reeve, 164 01-718 exusta Reeve, 127 01-653 falklandica Preston, 192 01-756 farquhari Turton, 106 (1-600 fasciata Krauss, 104 01-595 fasciolata Tomlin, 106 01-600 ferruginea Gmelin, 95 01-579 ficarazzensis de Gregorio, 96 01-580 flammmea Gmelin, 197 01-765 flava Hutton, 178 01-732 flemingi Powell, 186 01-740 flexuosa Quoy & Gaimard, 129 01-653 formosa Turton, 145 01-687 fragilis Philippi, 99 01-583 frauenfeldi Dunker, 98 01-582 fuegiensis Reeve, 197 01-765 fuenzalidai Herm, 142 01-676 fungoides Lightfoot, 125 01-651 fuscescens Gmelin, 11] 01-615 fuscoradiata Turton, 145 01-687 garconi Deshayes, 155 01-709 gemmula Turton, 145 01-687 gigantea Lesson, 142 01-676 gomesii Drouet, 100 01-584 [01-502] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Index 203 [looseleaf } [looseleaf ] gorgonica Lightfoot, 125 01-651 Laevipatella Pallary, 95 01-579 granatina Linnaeus, 109 01-613 laevis Pennant, 144 01-682 Granopatella Pallary, 95 01-579 lamarckii Payraudeau, 96 01-580 granularis Linnaeus, 115 01-629 laticostata Blainville, 135 01-659 grata Gould, 159 01-713 levata Deshayes, 128 01-654 guineensis Dunker, 82 01-558 limbata Philippi, 172 01-726 guttata d’Orbigny, 98 01-582 lineata Lamarck, 83 01-559 livescens Reeve, 154 01-708 longicosta Lamarck, 125 01-651 hamiltonensis Chapman & Gabriel oie ee ae 138 (1-662 uchuana Pilsbry, 87 56: hedleyi Oliver, 176 01-730 luctuosa Gould, 180 01-734 Helcion Montfort, 143 01-681 lugubris Gmelin, 112 01-616 Helcioniscus Dall, 147 01-701 lusitanica Gmelin, 96 01-580 helena Turton, 106 (1-600 luteola Lamarck, 96 01-580 helvola Turton, 106 (1-600 luzonica Reeve, 149 01-703 hentyi Chapman & Gabriel, 173 01-727 hepatica Verco, 137 01-661 hera Turton, 125 7 01-651 macquariensis Finlay, 199 01-767 hochstetteri K. Martin, 57 io 01-563 magellanica Gmelin, 198 01-766 hombroni Dautzenberg & Bouge, 153 OL-107 major Dautzenberg & Durochoux, 95 01-579 howensis Iredale, lid 01-728 major Pallary, 96 01-580 hyalina Philippi, 192 01-756 major Turton, 125 (1-651 margaritacea Gmelin, 99 01-583 margaritaria Reeve, 180 01-734 illuminata Gould, 185 01-739 maroccana Pallary, 98 01-582 imbricata Reeve, 178 01-732 mauritiana Pilsbry, 87 01-563 imperatoria de Gregorio, 96 01-580 maxima d’Orbigny, 142 01-676 inconspicuus: Suter, 150 01-734 mazatlandica Sowerby, 160 01-714 inquisitor Iredale, 130 01-654 melanostomus Pilsbry, 165 Ol=719 insignis Dunker, 158 01-712 meridionalis Rochebrune & Mabille, 198 — 01-766 intermedia Auct., 83 01-559 mestayerae Suter, 158 01-719 intermedia Bucquoy, Dautzenberg & ot metallica Rochebrune & Mabille, 195 01-766 Dollfus, 99 01-583 Phe ; e ee ee we . ‘ . ; — mexicana Broderip & Sowerby, 141 01-675 intermedia Jeffreys, 83 01-559 miliaris Turton, 116 01-630 intermedia Knapp, = 01-559 miniata Born, 118 01-636 2 os ad 01-979 minor Marion, 96 01-580 tere ala Purton, 126 Oi bes minor Steciel 108 (01-766 UnLOL tA Pennant, las 01-682 monopis Gmelin, 111 01-615 intraurea Iredale, 130 01-654 monopsis Turton, 126 01-652 morbida Reeve, 116 01-630 moreleti Drouet, 99 01-583 jacksoniensis Lesson, 170 01-724 moreli Deshayes, 128 01-654 jutsoni Chapman & Crespin, 82 01-558 multilirata Turton, 126 (1-652 mytiliformis Lightfoot, 192 01-756 mytilina Helbling, 192 01-756 kaffraria Rennie, 107 01-601 mytiloides Schumacher, 192 01-756 karachiensis Winckworth, 153 01-707 kerguelenensis E. A. Smith, 192 01-756 kermadecensis Pilsbry, 132 01-656 kowiensis Turton, 126 01-652 Nacella Schumacher, 191 01-755 kraussii Dunker, 113 01-617 Nacellinae Thiele, 147 01-701 [01-503] 204. Index A. W. B. Powell [looseleaf] natalensis Krauss, 116 01-630 neglecta Gray, 136 (1-660 nelsonensis Trechmann, 83 01-559 nigrisquamata Reeve, 161 01-715 nigrolineata Reeve, 161 01-715 nigro-punctata Reeve, 96 01-580 nigro-radiata Christiaens, 98 01-582 nigro-squamosa Dunker, 98 01-582 nigro-sulcata Reeve, 87 01-563 nimbus Reeve, 149 01-703 nodosa Hombron & Jacquinot, 180 01-734 novemradiata Quoy & Gaimard, 155 01-709 nympha Turton, 125 01-651 obtecta Krauss, 125 01-651 obtecta Turton, 127 01-653 octoradiata Hutton, 138 01-662 oculus Born, 110 01-614 oculus hirci Lightfoot, 111 01-615 Olana H. and A. Adams, 121 01-643 olivacea Hutton, 152 01-736 oliveri Powell, 187 01-741 opea Reeve, $7 01-563 optima Pilsbry, 131 01-655 orichalcea Philippi, 182 01-736 orientalis Pallary, 95 01-582 orientalis Pilsbry, 151 01-705 ornata Dillwyn, 179 01-733 ovalis Pilsbry, 125 01-651 pallida Gould, 87 01-563 Patella Linnaeus, 94 01-578 Patellanax Iredale, 123 01-649 Patellastra Monterosato, 95 01-579 Patellidae Rafinesque, 94 01-575 Patellidea Thiele, 115 01-629 Patellona Thiele, 109 01-613 Patellopsis Thiele, 95 01-579 Patellus Montfort, 95 01-579 patera Réding, 155 01-712 Patina Gray, 144 01-682 Patinastra Thiele, 145 01-687 Patinella Dall, 193 01-761 Patinigera Dall, 193 01-761 patriarcha Pilsbry, 127 01-653 paumotensis Gould, 130 01-654 pectinata Born, 144 01-682 pectinatus Montfort, 144 01-682 pectunculus Gmelin, 143 pellucida Linnaeus, 144 Penepatella Iredale, 123 pentagona Reeve, 130 perana Iredale, 182 percostata de Gregorio, 96 peronii Blainville, 136 perplexa Pilsbry, 138 petalata Reeve, 149 pholidota Lesson, 182 phymatias Suter, 183 pica Reeve, 128 picta Jeffreys, 95 pilsbryi Brazier, 133 piperata Gould, 98 planulata Turton, 111 plicaria Gmelin, 125 plicata Born, 125 plumbea Lamarck, 113 polaris Hombron & Jacquinot, 195 polygramma Tomlin, 106 pottsi Hutton, 189 pricei Powell, 155 profunda Deshayes, 87 prolixus Oliver, 176 pruinosa Krauss, 145 pulchella Turton, 119 pulchra Lightfoot, 119 punctata Lamarck, 96 pupillata Rochebrune & Mabille, 198 pyramidata Lamarck, 96 radians Gmelin, 180 radiata Born, 148 radiata Krauss, 104 radiata Perry, 95 radiatilis Hombron & Jacquinot, 182 rangiana Rochebrune, 107 redimiculum Reeve, 188 reevei Hutton, 178 reussi K. Martin, 83 reynardi Pilsbry, 149 reynaudi Deshayes, 149 rietensis Turton, 106 rosea Réding, 119 rota Gmelin, 149 rouxii Payraudeau, 96 rubicunda Réding, 119 rubraurantiaca Blainville, 172 rufanensis Turton, 145 rumphii Blainville, 158 [01-504] Patellidae [looseleaf] 01-681 01-682 01-649 01-654 01-736 01-580 01-660 01-662 01-703 01-736 01-737 01-654 01-579 01-657 01-582 01-615 01-651 01-651 01-617 01-763 01-600 01-743 01-709 01-563 01-730 01-657 01-637 01-637 01-580 01-766 01-580 01-734 01-702 01-595 01-579 01-736 01-601 01-742 01-732 01-559 01-703 01-703 01-600 01-637 01-703 01-580 01-637 01-726 01-687 01-712 November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 [looseleaf] rustica Linnaeus, 96 01-580 rustica Menke, 136 01-660 rustica Reeve, 127 01-653 safiana Lamarck, 113 01-617 sagittata Donovan, 158 01-712 sagittata Gould, 159 01-713 sandwichensis Pease, 165 01-719 sanguinans Reeve, 119 (1-637 sanguinolenta Gmelin, 119 01-637 scalata Reeve, 149 01-703 schroeteri Krauss, 111 01-615 scopulinus Oliver, 176 01-730 scutellaris Lamarck, 111 01-615 scutellaris: Reeve, 99 01-583 Scutellastra H. and A. Adams, 123 01-649 secernenda Dautzenberg, 95 01-579 silicina R6ding, 99 01-583 sitta de Gregorio, 96 01-580 solida Blainville, 171 01-725 sontica Iredale, 170 01-724 sowerbyi Turton, 125 01-651 spectabilis Dunker, 54 01-560 spinifera Lamarck, 125 01-651 squamata Réding, 96 01-580 squamifera Reeve, 137 01-661 squamosa Gmelin, Turton, 126 01-652 stearnsii Pilsbry, 160 01-714 stellaeformis Reeve, 130 01-654 stellata Bucquoy, Dautzenberg & Dollfus, 99 01-583 stellifera Gmelin, 182 01-736 stellularia Quoy & Gaimard, 183 01-737 strigatella Rochebrune & Mabille, 195 (1-763 strigilis Hombron & Jacquinot, 153 01-737 subgranularis Blainville, 96 01-580 subplana Potiez & Michaud, 99 01-583 taberna Powell, n. sp., 190 01-744 tabularis Krauss, 126 01-652 tahitensis Pease, 167 01-721 taitensis Réding, 166 01-720 talcosa Gould, 166 01-720 tara Prashad & Rao, 130 tarentina Lamarck, 99 tasmanica Tension Woods, 137 tella Bergh, 144 tenuilirata Carpenter, 163 terroris Filhol, 199 tessellata Hombron & Jacquinot, 153 testudinaria Linnaeus, 156 thetis Turton, 125 thomsoni Powell & Bartrum, 189 tincta Rochebrune & Mabille, 198 tomlini Turton, 116 toreuma Reeve, 162 tramoserica Holten, 170 travancorica Preston, 149 tuamotuensis Dautzenberg & Bouge, 130 tucopiana Powell, 134 turbator Iredale, 169 umbella Gmelin, 119 undato-lirata Reeve, 165 ustulata Reeve, 137 variabilis Krauss, 104 varicosa Reeve, 195 variegata Blainville, 170 variegata Reeve, 145 variegata Reeve (1842), 150 venosa Reeve, 198 vidua Reeve, 116 vitiensis Powell, 158 vitrea Philippi, 192 vulcanicus Oliver, 176 vulgata Linnaeus, 95 watsoni Christiaens, 98 whitechurchi Turton, 125 zebra: Reeve, 136 [01-505] Index 205 [looseleaf } 01-654 01-583 01-661 01-652 01-717 01-767 01-707 01-710 01-651 01-743 01-766 01-630 O1=716 01-724 01-703 01-654 01-658 01-723 01-637 01-719 01-661 01-598 01-763 01-724 01-702 01-704 01-766 01-630 01-712 01-756 01-730 01-979 01-582 01-651 01-660 206 Patellidae A.W. B. Powell Published by THE DEPARTMENT OF MOLLUSKS Delaware Museum of Natural History Greenville, Delaware 19807, U.S.A, [01-506] November 27, 1973 5G4,05 £4 Mo oh INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpidae = 207 THE FAMILY HARPIDAE OF THE WORLD by Haracp A. REHDER Division of Mollusks National Museum of Natural History Smithsonian Institution Washington, D. C. Introduction The family Harpidae includes as their most dominant element the harp shells of the tropical seas. These striking and colorful shells have long been favorites with collectors, although because Plate 183. Above: Harpa amouretta Roding, showing head region with the inhalent siphon and the left tentacle with the eye at the base. Oahu, Hawaii. X 2. Photo by Olive Schoen- of the paucity of species in the genus and the relative rarity of most of the species, they have not been as popular a group as the cowries, the cone shells, and the volutes. They have, however, been the subject of icono- graphic treatment by numerous authors: Kiener (1835), Reeve (1843), Chenu (1853), Kiister (1857), Sowerby (1860), Tryon (1883), and Max- well Smith (1948). The only really critical study to date of the genus is that of Sutor (1877). Hedley (1911) published a short note updating the nomenclature of the species. berg. Below: Harpa major Réding, showing autotomized pos- terior portion of the foot. Nuku Hiva, Marquesas. X 1. Photo by Otis Imboden, National Geographic Society. [20-601] 208 H. A. Rehder Family characters The Harpidae is one of the seven families com- prising the superfamily Volutacea (considering the Vasidae and Turbinellidae as distinct fam- ilies) of the order Neogastropoda, and is usually placed near the Olividae and Turbinellidae. As a matter of fact, the subfamily Harpidae was once placed in the family Olividae, undoubtedly be- cause of the similarities in the characters of the foot and propodium and the extent of the pari- etal callus. The shell is characterized by possessing a large body whorl marked by more or less strong ribs, with the spiral sculpture, if present, of secondary importance. The protoconch in this small family is notable for the variety shown in its form and in the number of its whorls. These nuclear differ- ences are used to separate the genera and sub- genera, and are mentioned in the key (p. 000) and in the diagnoses of the supraspecific groups. For representations of the various types of proto- conchs see our illustrations. The aperture varies from broadly to narrowly ovate; the parietal and columellar areas of the ventral side are covered by a callus of varying extent. The anterior siphonal sinus is well-marked, a siphonal canal is very short or absent, and there is generally a shallow posterior sinus at the junction of the outer lip and the parietal wall. An operculum is absent. The living species of Harpa have a shell vividly ornamented with reddish brown and_ various shades of pink in more or less intricate patterns. The shells of Austroharpa are brightly colored, with or without spots, or their color pattern may be restricted to scattered spots. According to Quoy and Gaimard (1832-35, vol. 2, p. 617), shells of the males are always narrower than those of females. The foot of the animal is very large and fleshy, often with a rough, rugose surface, and is con- Plate 184. Radular teeth of: 1, Harpa amouretta Roding. X 600 (after Peile, 1939); 2, Harpa amouretta Roding. (after Harpidae spicuously divided into two parts. The propodium is broadly arcuate in front, terminating laterally in points that extend well beyond the width of the hind portion of the foot, or metapodium; the anterior margin may be shallowly scalloped and is sometimes more or less furrowed or grooved vertically. The propodium is joined by a broad neck to the metapodium which is elongately lanceolate, tapering posteriorward to a_ point, with its margins also gently and undulatingly scalloped. The head, when the animal is active, is usually hidden beneath the anterior edge of the shell with only the tentacles and long inhalent siphon vis- ible. The siphon protrudes through the siphonal notch, and the tentacles, when the animal is viewed from above, emerge from each side of the siphon. The tentacles are slender, pointed, and bear the eyes at the anterior end of a lateral en- largement above the base of the tentacles. From near the base of the head on the right side arises in males the elongate, tapering penis. In a speci- men of Harpa amouretta from Nuku Hiva, Mar- quesas, I was able to see the rather long extruded part of the proboscis. In the Harpidae this organ is of the pleurembolic type, in which only the basal part is invaginable, withdrawing the distal portion into a sheath in the body cavity; this sheath is depicted by Quoy and Gaimard in one of their drawings. All exposed parts of the animal are usually vividly mottled and flecked with various shades of brown or red, from deep chestnut brown to pale brown or reddish brown, and sprinkled with spots of bright yellow. The tentacles and inhalent siphon are irregularly ringed with chestnut brown and also spotted with yellow. The penis either is sprinkled with very fine spots of reddish brown or is without spots. According to Quoy and Gai- mard the base of the foot is lightly spotted or Bergh, 1901); 3, Harpa major Roding. (after Troschel, 1866). [20-602] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpidae 209 flecked with pale reddish brown. In preserved animals these colors first turn a more or less scar- let red and then usually fade. The living animals and the anatomy of Harpa major and H. amouretta have been described and figured by Quoy and Gaimard (1832-35, vol. 2, pp. 611-620, pl. 42), and Bergh (1901) has pub- lished anatomical notes on H. ventricosa, H. ma- jor, H. doris, and H. amouretta. Earlier Reynaud had described the anatomy of either major or davidis, based on specimens from Ceylon (of which the shells are not described). His paper was read in 1829 but was published later (Reynaud, 1834); the figures in his paper are rather crude. The radula of Harpa is very small, and Quoy and Gaimard were unable to find it in their exami- nation of over twenty animals. Macdonald was apparently the first to discover the radula, which he said was “very minute compared with the whole bulk of the animal, as to appear quite rudi- mentary” (Macdonald, 1957, pp. 389-390). The radula was described and figured by Macdonald (1869, p. 116, pl. 13, fig. 14), and in the same year Troschel described that of H. major (Troschel and Thiele, 1866-93, p. 105, pl. 10, fig. 1). Tros- chel suggested the possibility that the radula teeth disappear in the adult stage, since the speci- men of which he examined the radula was a juve- nile. This probably accounts for Cooke’s categori- cal statement that there is no radula in the adults of Harpa (Cooke, 1895, pp. 216, 221). Peile has figured and discussed the radula of H. amouretta (Peile, 1939, pp. 271-272) as has Bergh (1901, p. 625, pl. 47, fig. 21). For copies of some of these radula drawings, see plate 184. Attempts to ex- tract the radula from a specimen of Harpa costata were finally successful, but in the process of staining and fixing, the radula became twisted on itself, and it was not possible to get a flattened Plate 185. Egg capsules of Harpa major Réding from Mot Tibarama, east coast of New Caledonia. 1, single egg capsule showing the “pore” at the top and 7 attachment scars. 2, view from top of cluster of egg capsules (after Risbec, 1932). section of the radula. I was able, however, to con- firm Peile’s statement that the mesocone of the rachidian projects forward as a fang-like tooth. It appears that on each side of the mesocone is a single small cusp. On either side of the rachidian is a broad, segmented membrane that may, as Peile suggests, be the remains of the bases of lat- eral teeth. Biology Most of the living species of the genus Harpa live in moderately shallow to rather deep water where they progress over the sand or hard bottom and burrow in the sand by means of their large and strong foot. Reynaud (1834, p. 35) says that they move by using their propodium as a point of attachment and pull the rest of their body forward. I have not observed this in the living specimens I have seen, and other writers have not mentioned this means of progression. Their mode of burrowing in the sand is de- scribed in a letter from Mrs. E. Couacaud of Port Louis, Mauritius. She observed Harpa major en- tering the sand with the shell perpendicular “until it was completely covered with sand. Then im- mediately by two or three successive movements I understood it had taken the horizontal position again, and then the tip of the siphon appeared from beneath the sand.” Chabouis and Chabouis, in a natural history textbook for French Polynesia (Chabouis and Chabouis, 1954), say that after H. major has buried itself in the sand the only trace is a small hole in the center of a low mound which reveals the location of the siphon. Plate 186. Egg mass of Harpa amouretta Roding from Batangas Province, Luzon Island, Philippines. Left fig. 11 capsules, about {0 mm. in width. Right fig., single capsule, showing pore at right, and broken attachment points at the left. Photo courtesy of R. Tucker Abbott. [20-603 | 210 H. A. Rehder An aspect of the behavior of species of the ge- nus Harpa that has attracted the attention of even the older naturalists is that of autotomizing (self- amputation) the posterior portion of the foot when disturbed. Rumphius as early as 1705 mentions this habit. This trait, found also in some other genera of marine and land snails that possess a large foot, has been discussed at length by Quoy and Gaimard (op. cit.), Reynaud (op. cit), and Jousseaume (1883). Crichton (1941, p. 330) de- scribes the autotomy of the hind portion of the foot of H. davidis, emphasizing the readiness with which this occurs. Several specimens of H. major dredged in the Marquesas severed the posterior portion of the foot (pl. 183), the severed ends of each part showing a clean edge (pers. obs. 1967). We did not notice any line of demarcation where the break occurred; Reynaud (op. cit.) states that there is such a line, but Quoy and Gaimard deny this. Stasek (1967) has published a paper on auto- tomy in general, in which he cites the observations on Harpa by Quoy and Gaimard and Reynaud. In a recent article by Hardy (1972), autotomy in H. ventricosa and amouretta is described and figured. None of the previous writers have published anything on the food of Harpa. Quoy and Gaimard state that they found nothing in the more than twenty stomachs of Harpa they examined, and be- lieved that their food must be tenuous and readily assimilable. The first information on the feeding habits seems to be that of Chabouis and Chabouis (op. cit.). As this book, published in 1954, is not readily available, I give here a free translation of the pertinent paragraph: “We have ascertained that the harp shell feeds on small crabs living in the sand, principally the box crab, Calappa hepatica, and the swimming crab, Neptunus [=Portunus| sanguinolentus. The anterior portion of the foot [propodium] holds the crab immobile on the substrate and the pos- terior portion [metapodium] glides underneath the crab, rolling itself around the imprisoned crab. At the same time the Harpa secretes a sticky fluid which combines with the sand to form a sort of a coating over the crab, asphyxiating it. We have seen a Harpa leave the sand while discarding the empty carapaces, probably after a meal.” This observation is confirmed by Mrs. Couacaud in a letter from Mauritius in which she wrote that she has seen Harpa major “lying on one side with the animal rolled like a ball around a small crab or shrimp which it was surely eating. I do not re- member having noticed this with Harpa minor, however.” Harpidae It is possible that the mucus or some other se- creted fluid contains a toxin that immobilizes or kills the crab on which the Harpa feeds. How the carapace and other hard parts of the crab are at- tacked by the snail in order to get at the soft parts is not known. The long proboscis is well suited to suck up the juices of the victim, the tiny radula possibly acting to reduce the larger pieces of flesh. Harpa has several rather conspicuous mucus glands in the body cavity which can secrete large quantities of mucus when the animal is disturbed, as we discovered when picking out living speci- mens in the dredge hauls we made in the Mar- quesas. Quoy and Gaimard comment on the diffi- culties that the coagulated mucus caused during their dissection and examination of Harpa major. The only literature reference to the eggs of Harpa that I have found is the note by Risbec on a female of Harpa major (he used the name Harpa nablium Mart.) that he found in New Caledonia depositing eggs (Risbec, 1932, p. 368). The snail when discovered had not yet completed the task, and with her foot covered the whole egg mass which was attached to a rock. The ootheca are lamellar, rather tough, roughly rectangular in shape, about 30 mm. long and 15 mm. high, and irregularly attached to each other. The opening is in the center of the upper edge, and each cap- sule contained about three to four thousand yel- lowish white eggs. In shape the individual egg capsules are rather like those of certain species of Busycon, and when viewed from above the serially arranged capsules remind one of a portion of a Turbinella or a Busycon egg mass (Plate 185). Dr. R. Tucker Abbott kindly examined and photographed for me the egg masses of Harpa amouretta Roding. The first egg mass (Plate 186) was laid by a female (typical shell form, Pl. 189, figs. 6-8) from Batangas Province, Luzon, Philip- pines, on March 25, 1965, and consisted of eleven lamellar, rather pliable, tough, rectangular oo- theca, about 40 mm. in width and 20 mm. high. At the top of each capsule is a lanceolate pore with a thin membrane covering it. The pore and the continuing slit on each side offers a potential opening of 20 mm. in length. The capsules have 5 to 7 points of attachment to each other. The shell length of the female is 46.8 mm.; its width is 28.0 mm. The second egg mass also consisted of eleven capsules, but they were more arching in shape and only 25 to 30 mm. in width. They were laid on June 12, 1966, on Panay Island, Philippines, by a female whose shell size is 44.5 x 26.5 mm. Approximately 3,000 to 4,000 eggs were in each capsule. The egg capsules were collected [20-604] Harpidae 211 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 November 27, 1973 Plate 187. Harpa from the Indo-Pacific. 700307); 6, Indian Ocean (Del. Mus. Nat. Hist. 12248). Figs. 1, 2, 3. Harpa costata (Linné, 1758). 1, Fouquet Id., Mau- ritius (ANSP 265220); 2, immature from Mahébourg reefs, Mauritius (Del. Mus. Nat. Hist. 12246); 3, La Morne, Mau- Figs. 7-10. Harpa harpa (Linné, 1758). 7 and §, Losap Id., Mortlocks, Caroline Ids. (Del. Mus. Nat. Hist. 37567); 9, Siasi, Sulu) Archipelago, Philippines (USNM_ 612453); 10, ritius (USNM 666304). Figs. 4, 5, 6. Harpa davidis Riding, 1798. 4, Rameshwaram, southern India (USNM 290884); 5, Indian Ocean (USNM Ryukyu Island (USNM _ 343465). [20-605] 212 H. A. Rehder Harpidae and preserved by the well-known collector, James E. Norton, now of Arizona. Reynault has recently published on the charac- ters and origins of the apyrene spermatozoa found in Harpa amouretta (Reynault, 1965). Classification Linné placed the two species of harp shells that he described in his genus Buccinum. The first to give these shells a distinct generic appellation was Roding, who in 1798 used the name Harpa, including in this genus six validly named species. Lamarck a year later independently created the genus Harpa for the same group. The harp shells were first placed in a distinct family by Bronn, who in 1849 proposed the family Harpina (Bronn, 1849, p. 469), including in this family also the genus Dolium. Early classification systems generally assigned the genus Harpa to the family Buccinidae (Gray, 1847, p. 138; Wood- ward, 1851, p. 116). H. and A. Adams (1853-54, p. 116) put Harpa in the subfamily Harpinae of the family Dactylidae (= Olividae), probably on the basis of the similarity in the nature of the large foot, parietal callus, and the absence or small size of the operculum. In this arrangement they were followed by Gray (1557, p. 26). Chenu was the first to use the family name Harpidae in the modern sense (Chenu, 1859, p. 204), and was followed in this usage by all later workers. The first person who appears to have published on the differences between the then known living and fossil species was Jousseaume, who in a short paper (Jousseaume, 1881) pointed out these dif- ferences and suggested using the name Cithara Klein for the recent species, and restricting the name Harpa Lamarck to the fossil species. As the type species of Harpa is the recent species H. harpa Linné, Cithara ‘Klein’ Jousseaume, 1881, falls into the synonymy of Harpa Roding. In 1883 Fischer (1880-87, p. 601) proposed Eocithara as a section of Harpa for the fossil spe- cies. He also included Silia Mayer, 1877, as a sub- genus of Harpa; this group, however, should probably be placed in the Volutidae, as it was put with a query by Cossmann as a subgenus in Cryp- tochorda, which he assigned to the family Harpi- dae (Cossmann, 1899, p. 78); the latter genus is now generally placed in the Volutidae. For many years Eocithara was used as a sub- genus of Harpa; Wenz (1943, pp. 1309-1311) has four subgenera under Harpa—Harpa s.s., Eoci- thara, Marwickara Laws, 1935, and Austroharpa Finlay, 1931. Most recent workers in New Zea- land and Australia, however, consider Austro- harpa and Eocithara as distinct genera, an ar- rangement I am following in this paper. Distribution The living species of Harpidae are found in the tropical and subtropical waters of the world, ab- sent only from the western Atlantic. Of the two groups found living today, the genus Austroharpa, subgenus Palamharpa is restricted to the warm-temperate waters of southern Aus- tralia, from the southernmost part of Queensland, around the southern coast, to the vicinity of Perth, Western Australia. All these species live in mod- erately deep water, in 20 to 100 fathoms, and are rather rare. The genus Harpa includes the brightly colored, strongly ribbed harp shells well-known in collec- tions. Of the ten recognized species eight are found in the Indo-Pacific region. Only two of these are widely distributed over much the entire re- gion; these are amouretta and major. Harpa harpa is almost as widely spread, but is rare in the Indian Ocean, and is not found in Polynesia, its range reaching only to the Gilbert Islands in Micronesia and Tonga in eastern Melanesia. Two species, ventricosa and costata, have a re- stricted distribution in the western Indian Ocean. H. davidis is found only in the eastern Indian Ocean, and H. articularis is restricted to the south- western Pacific Ocean. Harpa gracilis is a rare species found only in the coral atolls of the central Pacific, from the Ellice Islands to the Tuamotus and Clipperton Island. The remaining two species of Harpa are found outside of the Indo-Pacific: H. doris along the West African coast and on Ascension Island, and H. crenata in the Panamic province. Most of the species of Harpa are found in mod- erately deep water, and have been collected alive only by dredging, diving, or being found in fish traps. Only of amouretta have we definite knowl- edge of being found in shallow water and on reef flats; gracilis may also have a similar habitat. Paleontological History The earliest members of the family Harpidae belong to the genus Eocithara, of which eleven nominate species (one species has three subspe- cies) are known, and which has been found in Tertiary deposits from the Upper Paleocene to the Middle Miocene. The oldest species is Eoci- thara morgani Cossmann and Pissaro, 1909, from the Ranikot beds of the Upper Paleocene of Paki- stan. A relatively recently discovered specimen [20-606] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpidae 213 belonging to this genus in Lower Eocene beds of New Zealand is another bit of evidence that at this early period an arm of the Tethys Sea must have extended to New Zealand. We now know, there- fore, that by Middle Eocene times the genus Eoci- thara had spread in the east to New Zealand, and to the westward as far as Texas and California. Upper Eocene species are known from Burma and Mississippi. Regarding the phylogenetic origin of the Harpi- dae we are still in doubt as to a possible ancestral form, and to spend much time on this problem is to my mind an exercise in futility. Cox has outlined the situation very well when he wrote “Although the apparent great faunal gap between the Creta- ceous and the Tertiary has diminished consider- ably with increased knowledge, it may still be said that we know nothing of the ancestry of most of the new molluscan groups that appeared during the period of intense evolutionary activity which marked the transition from Mesozoic to Cainozoic times... each new genus... appears more or less suddenly, without any evidence of having evolved gradually from older forms . . the more we are forced to the conclusion how in- ‘ the more we know, complete the record is, and may always remain” (Cox, 1930, pp. 142-143). If we cannot determine the origins of the Harpi- dae we can make some conjectures as to the later history of the family. It seems very possible that the family, along with other warm-water genera “first evolved in the Indian Ocean and_ subse- quently migrated westwards” (Cox, loc. cit.). In Middle Eocene times the genus Eocithara is found in Texas and as far west as California where it is represented by a subspecies of E. mutica of the Paris Basin, as well as by a distinct species. Another species is found in the Upper Eocene of Mississippi. To the east of India the genus persisted longer than in Europe, where its last representative is found in the Lower Oligocene. In Java a species of Eocithara is found in the Miocene, and in the Marshall Islands a fragment assignable to this genus has been found in beds of Upper Miocene age. From eocitharid stock arose a species found in the Upper Eocene of New Zealand that has been placed in a distinct subgenus, Marwickara Laws, 1935. Another species, of quite different aspect but with a parallel origin, is found in the Middle Miocene of southern Australia; this is now placed in the subgenus Refluharpa Iredale, 1931. Meanwhile a distinct group was arising, prob- ably derived from some early eocitharid stock. This was the genus Harpa, characterized by a rela- tively smaller, broadly conical spire, a greater fusion and extension of the successive expanded upper portions of the outer lip and parietal callus over the preceding whorl, and a thinner, broader, not strongly marginate parietal and columellar callus that is more closely appressed over the ven- tral surface of the siphonal fasciole. One of the key differential characters is the multispiral, elevated-conical protoconch with peripherally keeled whorls found in Harpa; this contrasts with the paucispiral protoconch of Eocithara with its somewhat planate apex. Unfortunately the protoconch of most of the specimens of fossil species assigned to Harpa is more or less eroded and thus the characters are not clear. Other characters, however, observed in certain fossil species, such as the nature of the parietal callus and its extension over parts of the penultimate whorl, have induced me to place them in the genus Harpa rather than in Eocithara. The oldest recognized species of Harpa is H. myrmia Olsson of the Lower Oligocene of Peru. Other species are found in the Lower and Middle Miocene of France, Italy, the Caribbean, and Fiji, and in the Pliocene of Japan. From Miocene times onward species now living are represented in the fossil record. In the Oligocene and Lower Miocene times there arose in southern Australia a stock of small fossil species which are now all placed in the ge- nus Austroharpa. These are distinguished from both Eocithara and Harpa by their paucispiral, bulbous or dome-shaped protoconch. Their phylo- genetic origin is unknown, but some of the Middle Miocene and Pliocene species gave rise to the four recent species that now live in the warm-temper- ate waters of southern Australia. [20-607] 214 H. A. Rehder List of Recognized Taxa The daggers (+) preceding the taxa indicate fos- sil forms. Family Harpidae Bronn, 1849 Genus Eocithara Fischer, 1883 Subgenus Eocithara Fischer, 1883 tmorgani Cossmann and Pissaro, 1909. Upper Paleocene, Pakistan tmutica Lamarck, 1803. Type. Middle Eocene, France tsubsp. altavillensis Defrance, 1821. Middle Eocene, France . tsubsp. hilarionis Gregorio, 1880, Middle Eocene, northern Italy tsubsp. californica Vokes, 1937. Middle Eo- cene, California tclarki Vokes, 1937. Middle Eocene, California traricostata Risso, 1826. Middle or Upper Eo- cene, France telegans Deshayes, 1835. Upper Eocene, France tiacksonensis Harris, 1596. Upper Eocene, Mis- sissipp1 tbirmanica Vredenburg, 1923. Upper Eocene, Burma tsubmutica Orbigny, 1852. Lower Oligocene, France tnarica Vredenburg, 1925. Oligocene, Pakistan tbellardii Sacco, 1890. Oligocene, Italy tmuticaeformis Martin, 1916. Lower Miocene, Java Subgenus Marwickara Laws, 1935 twaihaoensis Laws, 1935. Type. Upper Eocene, New Zealand Subgenus Refluharpa Iredale, 1931 tlamellifera Tate, 1889. Type. Middle Miocene, southern Australia Genus Harpa Roding, 1798 harpa Linné, 1758. Type. Recent, Indo-Pacific ttosa Aoki, 1966. Lower Pliocene, Japan amouretta Roding, 1798. ¢Pliocene to Recent, Indo-Pacific Harpidae gracilis Broderip and Sowerby, 1829. Recent, Micronesia and Polynesia articularis Lamarck, 1822. Upper Miocene to Recent, southwestern Pacific major Roding, 1798. {Miocene to Recent, Indo- Pacific davidis Roding, 1798. Recent, eastern Indian Ocean ventricosa Lamarck, 1816. Recent, western In- dian Ocean costata Linné, 1758. Recent, islands of west- ern Indian Ocean doris R6ding, 1798. Recent, west coast of Af- rica; Ascension Island tbroichom Cossmann, 1899. Lower Miocene, France tiosephiniae Sacco, 1890. Middle Miocene, Italy tamericana Pilsbry, 1922. Middle Miocene, Car- ibbean crenata Swainson, 1822. Recent, eastern Paci- fic tmyrmia Olsson, 1931. Lower Oligocene, Peru Genus Austroharpa Finlay, 1931 Subgenus Austroharpa Finlay, 1931 tpulligera (Tate, 1889). Type. Middle Miocene, southern Australia Subgenus Palamharpa Iredale, 1931 exquisita (Iredale, 1931). Type. Recent, south- eastern Australia loisae Rehder, 1973. Recent, southern Western Australia punctata (Verco, 1896). Recent, South Aus- tralia wilsoni Rehder, 1973. Recent, southern West- em Australia tsulcosa (Tate, 1889). Middle Miocene, south- ern Australia ttatei Finlay, 1931. Pliocene, South Australia tspirata (Tate, 1889). Middle Miocene, South- ern Australia ttenuis (Tate, 1889). Lower to Middle Miocene, southern Australia tabbreviata (Tate, 1889). Middle Miocene, southern Australia ; tclathrata (Tate, 1889). Lower Miocene, south- ern Australia tpachycheila (Tate, 1894). Upper Oligocene, southern Australia tcassinoides (Tate, 1889). Pliocene, southern Australia [20-608] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpidae t tr * vt » 3 oy * Plate 158. Harpa from the Indo-Pacific. Figs. 1, 2. Harpa ventricosa Lamarck, 1816. I, Zanziar (USNM (USNM_ 681738); 6, Cebu, Philippines (USNM_ 612451); 597093); 2, Mauritius (USNM 666302). 7, southwest Pacific (Del. Mus. Nat. Hist. 1743). Figs. 3, 4. Harpa kajiyamai Rehder, new species. 3, paratype Figs. 8-11. Harpa major Réding, 1798. 8 and 11, typical form from the Philippines; 4, holotype, Philippines (Nat. Sci. from Cebu, Philippines (USNM 543683) and Okinawa, Mus. Tokyo 41450). Ryukyu Ids. (USNM 670380); 9 and 10, dark form or forma ligata Menke (Australian Mus. C. 73513) from Melanesia. Figs. 5-7. Harpa articularis Lamarck, 1822. 5 , off Taiwan [20-609] bo ou 216 H. A. Rehder Harpidae Doubtful and erroneously assigned species Harpa crescentensis Weaver and Palmer, 1922 Range—Middle Eocene of Washington (Crescent formation). Remarks—This small species, the holotype mea- suring only 6 mm. in length, does not appear to be a true Harpa. The attenuate anterior end of the body whorl (the authors in their original descrip- tion speak of the “canal moderately elongate”) is unlike that found in any species of Harpidae, as is the relatively broad, high spire. Vokes (1937, p. 12, pl. 2, fig. 9) figures a supposed topotype, somewhat larger (9.3 mm.) than the type, which shows the broad high spire but the specimen has a more typically harpid aperture, without any nar- rowing at the siphonal canal. Possibly two species are involved here. Synonymy— 1922 Harpa crescentensis Weaver and Palmer, Univ. Wash- ington Publ. Geology, vol. 1, p. 40, pl. 11, fig. 21 (Port Crescent, Clallam Co., Washington). 1937 Harpa? crescentensis Weaver and Palmer, Vokes, Jour. Paleont., vol. 11, p. 12, pl. 2, fig. 9. 1942 Harpa crescentensis Weaver and Palmer, Weaver, Univ. Washington Publ. Geology, vol. 5, p. 498, pl. 95, figs. 12, 13. Harpa dechordata White, 1888 Range—Paleocene of Brazil. Remarks—This species, with broad rugose folds on only the upper half of the body whorl, has been placed in Pseudoliva (Olividae) by Harris (1896, p. 154). Synonymy— 1888 Harpa dechordata White, Arch. Mus. Nac. Rio de Ja- neiro, vol. 7, p. 136, pl. 13, figs. 7, 8 (Maria Farinha, Pernambuco, Brazil). 1896 Harpa [Pseudoliva] dechordata White, Harris, Bull. Amer. Paleont., vol. 1, no. 4, p. 154. Harpa bellardii var. madachi Noszky, 1940 Range—Oligocene of Hungary. Remarks—tThis variety seems to me to have very little relationship with the Italian Eocithara bel- lardii Sacco, and probably represents a distinct species. However, the name is based on an inter- nal mold with adhering remains of the shell, is described only very briefly, and from the illustra- tion does not resemble any species in the Harpi- dae known to me. The ribs, very stout and few in number, do not appear to curve forward at the suture. Without a knowledge of the characters of the aperture and ventral side of the shell it is im- possible to discuss the affinities of this species, and even to be certain it is a member of this family. Synonymy— 1940 Harpa_ bellardii var. madachi Noszky, Ann. Hist.-Nat. Mus. Hungarici. Min.-Geol.-Paleont., vol. 33, p. 34, pl. 2, fig. 11 (near Budapest, Hungary). Eoharpa sinuosa Stephenson, 1955 Range—Upper Cretaceous of Mississippi and Missouri (Ow! Creek formation). Remarks—Proposed as a new genus in the fam- ily Harpidae, Eoharpa does not appear to belong in this family. The attenuation of the anterior end into a fairly long canal, the relatively high spire with fairly straight-sided whorls, and the presence of strong tubercles on the parietal and columellar callus are characters not found in any fossil or recent members in the family. Synonymy— 1955 Eoharpa sinuosa Stephenson. U. S. Geol. Survey Prof. Paper 274-E, pp. 131-132, p. 23, figs. 3-6. Harpa? soréensis Eames, 1952 Range—Eocene of Pakistan Remarks—From the illustration and description I doubt very much that this species belongs in the Harpidae. It might be assigned to the Buccinidae s. lat. Synonymy— 1952 Harpa? soriensis Eames, Phil. Trans. Royal Soc. Lon- don, Series B, no. 631, vol. 236, p. 106, pl. 4, fig. 91 (Zinda Pir section, Kohat District, Pakistan). Harpa trimmeri Fleming, 1828 Range—Lower Eocene of London, England. Remarks—According to information kindly fur- nished me by C. P. Nuttall and J. Cooper of the British Museum (Natural History), this species is very probably the volutid Athleta tricorona (J. Sowerby, 1840) from the London Clay, Lower Eocene. Synonymy— 1828 Harpa trimmeri Fleming, History of British Animals, p. 342 (probably London Clay at Brentford). Harpa neozelanica Suter, 1917 Range—Middle Eocene (Bortonian) of New Zea- land (Waihao). Remarks—Marwick (1934) suggests that Suter’s type represents a gerontic specimen of what Su- [20-610] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpidae 217 ter in the same paper described as Tudicla neozelanica. The latter species Marwick placed in his new genus Fascioplex (op. cit. p. 15) in the Fasciolariidae. Synonymy— 1917 Harpa (Eocithara) neozelanica Suter, New Zealand Geol. Survey, Palaeont. Bulletin No. 5, p. 43, pl. 5, fig. 11 (Waihao River, South Canterbury, New Zealead). 1934 Fascioplex neozelanica (Suter), Marwick, Proc. Malac. Soc. London, vol. 21, p. 16, pl. 1, fig. 8 (Harpa (Eocithara) neozelanica Suter listed as synonym). 1966 Fascioplex neozelanicus (Suter), Fleming, New Zealand Dept. Sci. and Industr. Res., Bull. 173, p. 316, pl. 109, fig. 1341. (Harpa neozelanica Suter listed as synonym in explanation of plate). Institutional Abbreviations The following abbreviations for institutions are used in this paper: AMN—American Museum of Natural History, New York. AMS—Australian Museum, Sydney. ANSP—Academy of Natural Sciences, Philadel- phia. AUCK—Auckland Institute and Museum, New Zealand BM—British Museum (Natural History), London. BPBM—Bernice P. Bishop Museum, Honolulu. CAS—Californmia Academy of Sciences, San Fran- Cisco. DMNH—Delaware Museum of Natural History, Greenville. MCZ—Museum of Comparative Zoology, Cam- bridge, Massachusetts. MHNG—Museum d'Histoire Naturelle, Geneva. MIHINP—Museum National d'Histoire Naturelle, Paris NMV—National Museum of Victoria, Melbourne. RNHL—Rijksmuseum van Natuurlijke Historie, Leiden. SAM—South Australian Museum, Adelaide. TAU—Tel Aviv University, Tel Aviv. USGS—U. S. Geological Survey, Washington. USNM—National Museum of Natural History, Washington. WAM—Western Australian Museum, Perth. 7MC— Zoological Museum, Copenhagen. Acknowledgments Iam grateful to many individuals for assistance received of all sorts and in varying degrees during the research and writing involved in this paper. In the following list I have attempted to name everyone to whom I am in any way indebted, and whose help is not acknowledged elsewhere. To any I may have left unnamed, by oversight, I herewith give my heartfelt thanks. For help given in allowing me to study col- lections in their care, forwarding museum speci- mens in their charge, and sending me needed information and photographs of types, I thank the following individuals: W. K. Emerson and W. E. Olds, Jr—AMNH; D. F. McMichael, W. F. Ponder, and P. H. Colman—AMS; R. Robert- son and V. O. Maes—ANSP; N. F. Tebble and J. E. Taylor—BM; A. E. Kay and Y. Kondo— BPBM; A. G. Smith—CAS; R. T. Abbott—DMNH; W. J. Clench and K. J. Boss—MCZ; E. Binder MHNG:: B. Salvat—MHNP; P. Dance—National Museum of Wales; C. O. van Regteren Altena —RNHL,; Al Barasch—TAU; B. R. Wilson and R. W. George—WAM; J. Knudsen—ZMC; W. O. Cemohorsky—Auckland Institute and Museum; P. M. Narang—Taraporevala Marine Biological Station, Bombay. To the following collectors I am indebted for the loan of specimens in their collections and for valuable information: Mrs. E. Couacaud, Port Louis, Mauritius; Mrs. S. T. Delaney, Santa Barbara, California; Mrs. M. C. Gniffiths, Lakes Entrance, Victoria; D. Hurrell, Port Lincoln, Vic- toria; T. Munyan, Atlantic City, N. J.. W. A. Tren- erry, Sydney, New South Wales. For information on important specimens, as well as sending me photographs of type speci- mens I am grateful to Alan Beu of the New Zealand Geological Survey, J. H. Macpherson and T. A. Darragh—NMV; H. M. Laws—SAM; N. H. Ludbrook, formerly of the Geological Survey of Victoria; L. M. March—WAM; the late G. Thorn- ley of Australia. Assisted by a grant from the National Science Foundation (Grant 24304,) I was able to visit the major museums in Europe and examine their col- lections of Harpa. During field work in the Mar- quesas Islands in 1967, carried out with the help of a grant from the National Geographic Society (Grant No. 624), numerous specimens of Harpa were collected. Finally, for advising me on numerous questions and problems brought to them, and for help in many ways I am indebted to the following col- leagues at the National Museum of Natural His- tory—H. S. Ladd, J. P. E. Morrison, J. Rosewater, and W. P. Woodring. [20-615] 218 H. A. Rehder Harpidae Key to the genera and subgenera of the family Harpidae A Protoconch multispiral (3 to 5 whorls), elevated-conical, whorls with a peripheral keel AA Protoconch paucispiral (2% to 3 whorls), ee ere er errr er ee reeemee re e Harpa broadly conical or bulbous, whorls without joerg) 5) ive cal cic] (meee nena ere ee ree rr eter rn ere Vereen errerere ee crore erry an ener B B Protoconch mamillate with planate apex, and 24 to 3 convex whorls, suture impressed dase Daarahea ig cdc exe dasha ated amsattiog react aus gape annie GIR IROL eee eee eee ee C BB Protoconch bulbous or dome-shaped, with 2 or fewer adpressed whorls ..................... D C Aperture length less than 60% of total lenathy: sesic, ccsacsccssesedsconsacwends cgantediguesouss Marwickara CC Aperture length more than 702 of total length: oisciccesaanvtsendrsix asnncnteealetnisiealoiadsaletataees E D_ Protoconch bulbous, with large, inflated initial whorl... eee Austroharpa DD Protoconch dome-shaped, with small initial whorl] «0.0.0.0... eee cesta Palamharpa EK Axial ribs widely separated, less than 18, parietal callus strong with marginate edge; protoconch small, rather turrited ............ bras Gudtoa ia tcae tecend nese ee aieetaaeiccate Eocithara EE Axial ribs crowded, more than 35; parietal callus thin, without sharp edge; protoconch large, broader than high ...................008 sopniaet- bases staauomen caetiseany cake acne eaiawaate Refluharpa Plate 189, Harpa from the Eastern Pacific, Indo-Pacific and Eastern Atlantic (Opposite page) Figs. 1, 2. Harpa crenata Swainson, 1522. 1, Mazatlan, Sonora, Mexico (ANSP 250671); 2, Mulege Bay, Baja California, Mexico (USNM 12509). Figs. 3-5. Harpa gracilis Broderip and Sowerby, 1829. 3, Vahitahi, Tuamotu Ids. (USNM 613243); 4, Raroia, Tuamotu Ids. (USNM 698318); 5, Papeete, Tahiti (Del. Mus. Nat. Hist. 12830). Figs. 6-8. Harpa amouretta Rodding, 1798, typical form. 6, 7 Siasi, Sulu Archipelago, Philippines (Del. Mus. Nat. Hist. 5185); 8, Ryukyu Ids. (USNM 670470). Figs. 9-11. Harpa amouretta Réding, 1798, forma crassa Krauss, 1848. 9, Zanzibar (USNM 597117); 10, Cebu Id., Philippines (Del. Mus. Nat. Hist. 12828). 11, Mogadiscio, Somalia (USNM 673593). Figs. 12-14. Harpa doris Réding, 1798. 12 and 13, Senegal (ANSP 180950); 14, Cape Verde Islands (Del. Mus. Nat. Hist. 122249). Figs. 15, 16. Harpa doris Réding, 1798. Broad, heavy form. Ascension Island, South Atlantic. 15, (MCZ 278591); 16, (MCZ 232221), [20-616] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpidae 219 Plate 189, Harpa from the Eastern Pacific, Indo-Pacific and Eastern Atlantic (explanation on opposite page) [20-617] 220 H. A. Rehder Harpidae Selected Bibliography Adams, F. Another live Harpa amouretta. Hawaiian Shell News, vol. 14, no. 13, pp. 2, 5, fig. Adams, Henry and Adams, Arthur. 1853-54. The Genera of Recent Mollusca, vol. 1, London. x1+484 pp. Anonymous. 1961. The Molluscan family Harpidae. Hawaiian Shell News, vol. 9, no. 6, p. 8. Anonymous. 1972. Le belle conchiglie da collezionione. La Conchiglia, col. 4, pp. 5-9. Bergh, R. 1901. Beitrag zur Kenntniss der Gattung Harpa. Zool. Jahrb. Abth. Anat. Ont. Thiere, vol. 14, pp. 609-629, pl. 47. Bronn, Heinrich G. 1849. Handbuch der Geschichte der Natur, vol. 3, pt. 3, Index Palaeontologicus. Stuttgart. 976 pp. Buchanan, J. B. 1954. Marine Molluscs of the Gold Coast. Jour. West African Sci. Assoc., vol. 1, pp. 30-45, 7 figs. Carpenter, Walter N. 1961. Ten Harpa costata. Hawaiian Shell News, vol. 10, no. 2, p. 4. Chabouis L. et Chabouis F. 1954. Petite Histoire Naturelle des Etablissements Frangais de VYOcéanie. Saint-Amand-Mon- trond, Cher. Chenu, J. C. 1853. Genus Harpa. Illustrations Conchylio- logiques, vol. 4 (pt. 85), 4 pls. Chenu, J. C. 1859. Manuel de Conchyliologie et de Paléontol- ogie Conchyliologique, vol. 1, Paris. viit508 pp., 3707 text- figs. Cooke, A. H. 1895. Molluscs, in the Cambridge Natural His- tory, vol. 3, xii + 459 pp., 311 text-figs. Cossmann, M. 1899. Essais de Paléoconchologie comparée, vol. 3, 201 pp., 8 pls., 34 text-figs. Cotton, Bernard C. and Woods, Nelly Hooper. 1933. Remarks on the new Harpid (Mollusca) genera of Finlay and Iredale. Records South Australian Museum, vol. 5, pp. 45-47, 9 figs. Cox, L. R. 1930. The Fossil Fauna of the Samana Range and some neighboring areas: Part VII. The Mollusca of the Hangu Shales. Mem. Geol. Survey India. Paleont. Indica, Crichton, M. D. 1941. Marine shells of Madras. Jour. Bombay Nat. Hist. Soc., vol. 42, pp. 323-341, 4 pls. Dance, S. Peter. 1966. Shell collecting. An Illustrated History. Berkeley and Los Angeles. 344 pp., 35 pls., 31 text-figs. Dance, S. Peter. 1967. Report on the Linnaean shell collection. Proc. Linn. Soc. London, vol. 178, pp. 1-24, pls. 1-10. Finlay, H. J. 1931. On Austrosassia, Austroharpa, and Austro- lithes, new yvenera; with remarks on the gastropod proto- conch. Trans. New Zealand Inst., vol. 62, pp. 7-19 (May 31). Fischer, Paul. 1880-87. Manuel de Conchyliologie et de Palé- ontologie Conchyliologique. Paris. xxiv + 1369 pp., 23 pls., 1138 text-figs. Garrard, T. A. 1961. Mollusca collected by M. V. “Challenge” off the east coast of Australia. Jour Malac. Soc. Australia, no. 5, pp. 2-37, 1 pl., 1 map. Gray, John E. 1847. A List of the Genera of Recent Mollusca, their Synonyma and Types. Proc. Zool. Soc. London, pt. 15, pp. 129-219. Gray, John E. 1857. Guide to the systematic distribution of Mollusca in the British Musuem. Part 1. London xii + 230 pp. Habe, Tadashige. 1961. Coloured Illustrations of the Shells of Japan. (II). Osaka. ix+183 pp., 64 pls., text-figs. [in Japanese]. Habe, Tadashige. 1964. Shells of the Western Pacific in Color, vol. 2. Osaka. 233 pp., 66 pls., text-figs. Hanley Sylvanus. 1855. Ipsa Linnaei Conchylia. London. 556 pp., 5 pls. Harrison, Ibby. 1968. Two rare Hawaiian shells live collected. Hawaiian Shell News, vol. 16, no. 7, p. 1, 4 figs. Hardy, R. 1972. L’ “autotomia” in Harpa major (= ventricosa). La Conchiglia, vol. 4, pp. 4, 6, 3 text-figs. Hedley, Charles, 1899. The Mollusca of Funafuti, in The Atoll of Funafuti, etc. Australian Museum Mem. III, pp. 395-510, 549,565, 80 text-figs. Hedley, Charles. 1911. The nomenclature of Harpa. Nautilus, vol. 25, pp. 65-66. Herrmannsen, A. N. 1846-47. Indicis Generum Malacazoorum Primordia, Cassel. vol. 1, xxviit+637 pp. Hertlein, Leo G. 1957. Pliocene and Pleistocene fossils from the southern portion of the Gulf of California. Bull. S. California Acad. Sci., vol. 56, pp. 57-75, 1 pl. Jacobs, George E. 1961. A review of the molluscan family Harpidae. New York Shell Club Notes, no. 68, pp. 3-4. Janus, H. 1961. Die Typen und Typoide stidafrikanischer Meeresmollusken im Staatlichen Museum ftir Naturkunde in Stuttgart. I. Gastropoda. Stuttgarter Betrige zur Natur- kunde, no. 70, 19 pp., 4 pls. Jousseaume, F. 1881. Observations relatives aux Mollusques du genre Harpa. Bull. Soc. Zool. France, vol. 5: Extr. Proc.-Verb., pp. XXXVIU-XXXVIII. Jousseaume, F. 1883. De animal d'une Cithara d’aprés d'une observation de M. A. Marche. Bull. Soc. Zool. France, vol. 8, pp. 205-208. Kiener, L. C. 1835. Genre Harpe. Spécies Général et Icono- graphie des Coquilles Vivantes. Paris, 12 pp. 6 pls. Kiister, H. C. 1857. Die Gattungen Cassis, Cassidaria, Oni- scia. Dolium, Eburna und Harpa. Syst. Conchylien-Cabinet, vol. 3, pt. 1B, 104 pp. pls. 36-70. Ladd, Harry S. 1966. Chitons and Gastropods (Haliotidae through Adeorbidae) from the Western Pacific Islands. U. S. Geol. Survey Prof. Paper 531, iv+98 pp., 16 pls., 14 textfigs., 6 tables. Macdonald, John D. 1857. Observations on the natural af- finities and classification of gasteropoda. Proc. Royal Soc. London, vol. 8, pp. 354-393. Macdonald, John D. 1869. On the Homologies of the Dental Plates and Teeth of Proboscidiferous Gasteropoda. Ann. Mag. Nat. Hist., ser. 4, vol. 3, pp. 113-117, pl. 13. Martin, K. 1879-80. Die Tertiirschichten auf Java nach den Entdeckungen von Fr. Junghuhn. Leiden, ixtl64+iv+51+3 pp., 28 pls. 1 map. Oostingh, C. H. 1938. Die Mollusken des Pliocans von Sud- Bantam in Java. I Gastropoda I. De Ingenieur in Neder- landsch-Indie, IV: Mynbouwen Geologie, vol. 5, pp. 17-33, 35-46, 49-60, 105-115, 7 pls. Palmer, R. H., and Hertlein, L. G. 1936. Marine Pleistocene Mollusks from Oaxaca, Mexico. Bull. S$. Calif. Acad. Sci., vol. 35, pp. 65-81, pls. 18, 19. Parker, R. H. 1964. Zoogeography and ecology of some macro-invertebrates, particularly mollusks, in the Gulf of California and the continental slope off Mexico. Vidensk. Medd. fra Dansk naturh Foren., vol. 126, 178 pp., 15 pls. Quoy, J. R. C., and Gaimard, J. P. 1832-35. Voyage de. . . lAstrolabe . . . pendant . . . 1826-29, sous le commande- ment de M. J. Dumont d'Urville, etc. Zoologie, vols. 2-3. 686+954 pp., 93 pls. Reeve, Lovell. 1843. Monograph of the genus Harpa. Conch- ologia Iconica, vol. 1, 4 pls., 6 pp. Reynaud, A. 1834. Observations sur l’animal de la Harpe. Mém. Soc. Hist. Nat. Paris, vol. 5, pp. 33-40, pl. 3. Reynault, Liliane. 1965. Origine et caractéres des spermato- zoides apyrénes de Harpa minor Lmk. (Mollusque Gastéro- pode Prosobranche). Comptes Rendus hebdom. Séances Acad. Sci. (Paris). vol. 260, pp. 665-667, 1 pl. Risbec, J. 1932. Notes sur la ponte et le developpement de mollusques gastéropodes de Nouvelle-Calédonie. Bull. Soc. Zool. France, vol. 57, pp. 358-374, figs. Rosewater, Joseph. 1968. Itinerary of the Voyage of H. M. S. Blossom, 1825 to 1828. Veliger, vol. 10, pp. 350-352. Salvat, B. and Ehrhardt, J. P. 1970. Mollusques de l'Tle Clipperton. Bull. Mus. Nat. d’Hist. Nat. Paris, ser. 2, vol. 42, pp. 223-231. [20-618] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpidae 221 Schepman, M. M. 1907. Mollusken aus posttertiaren Schichten von Celebes. Samml. Geol. Reichsmus. Leiden, S. I., vol. 8, pts. 3, 4, pp. 151-203, pls. 10-13. Sherborn, Charles D. 1940. Where is the —— Collection? Cambridge. 148 pp. Smith, Maxwell. 1948. Triton Helmet and Harp Shells. Winter Park, Fla. v+57 pp., 16 pls. Sowerby, G. B., I. 1860. Monograph of the genus Harpa. Thesaurus Conchyliorum, vol. 3, pp. 169-172, pls. 231-233. Stasek, Charles R. 1967. Autotomy in the Mollusca. Occ. Papers Calif. Acad. Sci. no. 61, 44 pp., 11 text-figs. Sutor Aug. 1877. Das Genus Harpa. Eine conchyliologische Studie. Jahrb. deutsch. Malakozool. Ges., vol. 4, pp. 97- 129, pls. 4, 5. Troschel, F. H. (and Thiele, J.) 1866-93. Das Gebiss der Schnecken zur Begriindung einer naturlichen Classification, vol. 2. Berlin, viiit409 pp., 32 pls. Tryon, George W., Jr. 1883. Subfamily Harpinae. Manual of Conchology, vol. 5, pp. 97-100, pls. 40-41. Verco, Joseph C. 1913. Note on Harpa (Eocithara) punctata Verco. Trans. Royal Soc. South Australia, vol. 37, pp. 446-447. [20-619] Vokes, H. E. 1937. The gastropod genus Harpa in the Eocene of the western United States. Jour. Paleont., vol. 11, pp. 10-12, pl. 2, figs. 1-8. Wagner, Robert J. L. and Abbott, R. Tucker. Van Nostrand’s Standard Catalog of Shells. Ed. 2. Princeton xit+303 pp., illustr. Weaver, Clifton S. 1963. Harp Shells in Hawaii. Hawaiian Shell News, vol. 11, no. 12, p. 1, 5 figs. Weaver, Clifton S. 1966. Common Pacific Harp rare in Hawaii. Hawaiian Shell News, vol. 14, no. 8, p. 8, figs. Wenz, W. 1943. Gastropoda, Pt. 6: Prosobranchia, in Schin- dewolf, Handbuch der Palaozoologie, vol. 6. Berlin, pp. 1201-1505, text-figs. 3417-4211. Woodward, S. P. 1851. A Manual of the Mollusca; or a Rudimentary Treatise of Recent and Fossil Shells. London. vilit 158 pp., 12 pls., 89 text-figs. Zwierzycki, J. 1915. Voorloopig Onderzoek van fossilien afkomstig van eenige Vindplaatsen op Sumatra. Jahrb. Mijnwezen Ned. Ost-Indie, vol. 42 (1913), pp. 101-129, pl. 3. bo bo to H. A. Rehder Harpidae bait eau on Harald A. Rehder, a well-known American mala- cologist, is now Senior Zoologist in the Depart- ment of Invertebrate Zoology, in the National Museum of Natural History, Smithsonian Insti- tution, Washington, D. C. He has his office in the Division of Mollusks, where he has worked for over forty years, having been curator of the Division from 1946 to 1966. Dr. Rehder was born in Boston, Massachusetts, on June 5, 1907, and received his A.B. at Bowdoin College, Brunswick, Maine, his M.S. at Harvard Univer- sity, and his Ph.D. at The George Washington University. For many years he has concentrated his research efforts on the marine mollusks of the Indo-Pacific region, particularly those of Polynesia. In connection with this research pro- ject he has made eight trips to the tropical Pacific in the last seventeen years, on the last two of which his wife, Lois, has accompanied him. Since 1924, he has published numerous scientific reports on mollusks, particularly those of the Caribbean and Hawaii. The Rehders have two grown children. [20-620] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Eocithara — 223 Family Harpidae Bronn, 1849 Genus Eocithara Fischer, 1883 This extinct genus comprises a group of species known only from the Lower Eocene to the Upper Miocene. They are all small, reaching a maxi- mum length of 39 mm. (about 1% inches), and have a paucispiral protoconch that is bluntly mammillate to broadly conical, with convex whorls and impressed sutures. The axial ribs or varices may be few (about twelve) or numerous and crowded and may be lamellar and some- times slightly reflected or ridgelike and triangu- lar in cross-section; the ribs at the shoulder are without spines, angular, or occasionally weakly spinose. The genus Eocithara sensu lato comprises three subgenera: Eocithara, sensu stricto, Marwickara Laws, 1935 and Refluharpa Iredale, 1931. These subgenera have been summarily differentiated in the key given above. The basic character differentiating Eocithara from the other genera of Harpidae is the pauci- spiral protoconch with a planate apex, and with the nuclear whorls without a peripheral keel but with impressed sutures. From Harpa it differs also in the generally much smaller size of its species, in the forwardly expanded upper part of the successive outer lips of the body whorl not fusing and covering most of the preceding whorl, and in the outer edge of the parietal callus being distinctly marginate. The genus Austroharpa differs in possessing a paucispiral, bulbous or dome-shaped protoconch, and in the forward curving upper part of the varices not prominent, or only slightly visible at the suture. Subgenus Eocithara Fischer, 1883 Type: Harpa mutica Lamarck, 1803 The members of the typical subgenus are marked by their rather broadly ovate shape, short conical spire, well-developed parietal callus that is margined at its outer edge often leaving exposed a pseudumbilical chink at the siphonal fasciole. The ten described species and subspecies as- signed to FEocithara s.s. occur in deposits of Lower Eocene to Lower Miocene age from Cali- fornia’ and Mississippi through France and Northern Italy to Burma, Java, and Fiji. A frag- ment apparently assignable to this subgenus was found in deposits of Upper Miocene age in the Marshall Islands. Synonymy— 1881 Harpa Lamarck, Jousseaume, Bull. Soc. Zool. France, vol. 5: Proc.-Verb, p. XXXVIII. Not Harpa Lamarck 1799, nor Harpa Roding, 1798. 1883 Eocithara Fischer, Man. de Conch., p. 601 (type by monotypy: Harpa mutica Lamarck.) Eocithara morgani (Cossmann and Pissaro, 1909) (Pl. 191) Range—Upper Paleocene of Pakistan (Upper Ranikot). Remarks—This, the earliest known species un- questionably assigned to the family, is un- doubtedly closely related to E. mutica Lamarck from the Middle Eocene of France, in the ances- tral line of which E. morgani may be considered to stand. A copy of the original description fol- lows: “Size moderate, shape depressed; spire conical, terminated by a smooth papillate protoconch of one whorl and a half; seven spire whorls, con- vex, depressed, separated by deep sutures, or- namented with thin, lamellar, distant ribs, slightly spinose posteriorly, regularly coinciding in suc- cessive whorls, decussated on the earlier whorls Plate 190. Eocithara mutica (Lamarck). Protoconch of speci- men from Chaussy, France; Middle Eocene. USNM _ 646907 X 10. [20-623] 224 ~=Eocithara H. A. Rehder Harpidae Plate 191. Eocithara morgani (Cossmann & Pissaro, 1909). Lower Eocene of Pakistan. Left-hand figure 33.9 mm. in length; right-hand figure 19.6 mm. (from Cossman & Pissaro, 1909, pl. 2, fig. 25; pl. 3, fig. 24). only, by a few spiral threads which are so thin as to be scarcely visible. Body-whorl very large, constituting almost the entire shell, excavated at its base. The axial lamellae persist upon the body-whorl up to its anterior region, while two or three axial threads, very thin, and of very slight relief, are intercalated amongst them, without spiral striations. It is only towards the base, in the excavated portion neighbouring the neck, that one distinguished some very fine spiral threads.” A plaster cast of the holotype, forwarded to me by M. V. A. Sastry of the Geological Survey of India, measures 33.9 mm. in length and 19.0 mm. in width. The type locality is given by Cossmann and Pissaro as north of Leilan coal pit, Sind, Paki- stan, from the upper beds of zone 3. Vredenburg (1928, p. 35) states that this is erroneous, and Plate 192. Eocithara mutica (Lamarck, 1807) Middle Eocene, Chaussy, France. USNM 646907 29.7 mm. [20-624] that the two syntypes are from the uppermost beds (zone 4) of the Ranikot at Jhirak, Sind. Vredenburg (1928, pp. 33-35) gives a more extended and detailed description of this species. Synonymy— 1909 Harpa morgani Cossmann and Pissaro, Mem. Geol. Survey India, Paleontologia Indica, n. s., vol. 3, mem. 1, p. 22, pl. 2, fig. 25, pl. 3, fig. 24. 1923 Harpa (Eocithara) morgani Cossmann and _ Pissaro, Vredenburg, Rec. Geol. Survey India, Paleontologia Indica, vol. 10, mem. 4, pp. 33-35 (Jhirak, Sind, Pakistan, Upper Ranikot). Eocithara mutica (Lamarck, 1803) (Pl. 192) Range—Middle Eocene of France (Lutetian of Paris Basin). Remarks—The type-species of Eocithara is a rather distinctive shell, and although not common in collections is the best known of all fossil species of the family Harpidae. The following description is based on five specimens in the collections of the National Museum of Natural History. The species that Deshayes depicts for Harpa mutica Lamarck (Deshayes, 1824, pl. 86, figs. 14, 15) represents his “Var. a” (Deshayes, idem, p. 642), which he describes as being larger, more inflated, and with more distant ribs. This is quite distinct from the typical mutica Lamarck, and may be the E. raricostata Risso discussed below. Description—Adult shell rather small, 24 to 39 mm. (1 to 1% inches) in length, ovate, with a coni- cal spire. Protoconch of 24 to 2% whorls, with flattened apex and impresséd suture, the whorls convex and smooth. Postnuclear whorls of spire convex, with sharp, lamellar, rather widely sepa- rated axial ribs that curve forward in the upper third of their length; in the body whorl this for- ward-curved upper end of each rib is adnate to the suture but does not reach the succeeding rib. Be- tween the axial ribs there are strong, somewhat distant, spiral striae, subequal in strength, and crossed by distant, axial growth lines; below the suture, on the subsutural ramp, the spiral lines are fewer and more distantly spaced. The body whorl is gently convex, occasionally showing a distinct angulation at the shoulder, in which cases the spiral sculpture on the subsutural ramp is ob- scure or absent; axial ribs 12 to 14 in number, lamellar, occasionally slightly reflected, especially near the aperture, at the shoulder sometimes no- ticeably angulate; in the intercostal spaces the distantly spaced growth wrinkles are prominent, crossing the strong spiral sculpture and giving the November 27, 1973 surface a netted appearance; siphonal fasciole strong, crossed by the strongly lamellose anterior end of the axial ribs. Aperture narrowly ovate, outer lip gently curved, smooth, reflected; inner lip forming a narrow callus on the parietal and columellar areas, with a conspicuous, slightly thickened margin, that in its anterior half is rather erect, and leaves exposed the pseudumbilical chink. Measurements (mm.)— length width no. whorls 29.7 18.0 6.5 Chaussy 39.0 29.0(?) original description 39.0° 24.9° Chaussy °Measured from figure in Cossmann and Pissaro, 1913 Synonymy— 1803 Harpa mutica Lamarck, Ann. Mus. Hist. Nat. Paris, vol. 2, pt. 2, p. 167; 1805, Lamarck, Ann. Mus. Hist. Nat. Paris, vol. 6, p. 227, pl. 44, fig. 14; 1824, De- shayes, Deser. Coq. Foss. Eny. Paris, vol. 2, p. 642 [not pl. 86, figs. 14, 15 which represent Deshayes’ var. a]; (Grignon, France). 1883 Harpa (Eocithara) mutica) Lamarck, Fischer, Manuel Conch., p. 601; 1899, Cossmann, Essais Paléoconch. Comp., pt. 3, pp. 73, 75, fig. 10, pl. 3, figs. 22, 23; 1913, Cossmann and Pissaro, Icon. Compl. Coq. Foss. Eocene Paris, vol. 2, pl. 46, figs. 209-1. Eocithara mutica altavillensis (Defrance, 1821) Range—Middle Eocene of France (Calcaire Grossier of western France). Remarks—This form, described as a species by Defrance, has been declared to be merely a variety of Eocithara mutica (Lamarck) that is smooth be- tween the ribs (Deshayes, 1865, p. 524). Without Plate 193. Eocithara’ mutica californiensis (Vokes, 1937). Middle Eocene of California. 26.4 mm. (from Vokes, 1937, pl. 2, figs. 2, 6). INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Eocithara = 225 seeing a specimen or figure, or even a description of this form I am unable to determine whether it should be ranked as a subspecies or a distinct species; for the present, therefore I am leaving it as a subspecies. Synonymy— 1821 Harpa altavillensis Detrance, Dict. Sciences Naturelles, vol. 20, p. 303 (Hauteville, Dept. de la Manche). 1865 Harpa mutica var. altavillensis Defrance, Deshayes, Descr. Anim. sans Vert Bassin de Paris, vol. 3, p. 524. Eocithara mutica californiensis (Vokes, 1937) (PI. 193) Range—Middle Eocene of California (Domen- gine stage). Remarks—V okes is undoubtedly correct in con- sidering this a subspecies of the Harpa mutica Lamarck from the Calcaire Grossier of France. The differences mentioned by Vokes are slight— one less axial rib on the body whorl and the pres- ence of spiral sculpture on the subsutural ramp. Description (copied from Vokes, 1937)—Types: Holotype, Univ. California Mus. Paleont. 30438; paratypes, U.C.M.P. 30439, 30440; loc. 3296, Llajas formation. Aliso Canyon, Ventura County, California. Dimensions: Holotype, length 26.4 mm., diam- eter (crushed), 16.2 mm.; paratype 30439, length, 19.8 mm., diameter (crushed), 9.7 mm.; paratype 30440, length (incomplete), 18.2 mm., diameter, 11.2 mm. Shell of moderate size, subfusiform, globose; nucleus smooth, papillate, of approximately three whorls; post-nuclear whorls five, convex, shoul- dered, with somewhat appressed sutures, orna- mentation consisting of prominent, distant, lamel- lar axial ribs decreasing in number from 15 to 17 on the first post-nuclear whorl to 10 to 12 on the body whorl; interspaces ornamented with numer- ous (6 to 15), secondary axial riblets; spiral sculp- ture of numerous, irregularly spaced primary and secondary riblets, the secondary ribbing tending to extend up on the shoulder of the whorl; aperture more than twice the length of the spire, moderate- ly narrow; outer lip simple, inner lip with a well- developed characteristic callus wash, anterior canal short; base of shell concave, with a well- developed fasciole. Synonymy— 1936 Harpa (Eocithara) mutica n. subsp. Vokes mss., Bull. Geol. Soc. America, vol. 47, p. 871 [nomen nudum. ] 1937 Harpa (Eocithara) mutica californiensis Vokes, Jour. Paleont., vol. 11, p. 11, pl. 2, figs. 2, 4, 6, 8. [20-625] 226 ~~ Eocithara H. A. Rehder Harpidae Eocithara mutica hilarionis (Gregorio, 1880) (Pl. 194) Range—Middle Eocene of Northern Italy (Upper Lutetian). Remarks—This subspecies is very close to the nominal species, and its validity as a distinct taxon is somewhat doubtful. The principal dif- ferences according to the author are the smaller number of ribs—10 or 11 rather than the 12 to 14 found in mutica s. str.—and stronger cancellate sculpture between the ribs. Because Gregorio’s work is rare and unavail- able to most students, I give a free translation of the original diagnosis: “It is rather common at San Giovanni, although complete specimens are rarely found. It reaches there a fair size: the figured specimen has a length of 35 mm., a width of 23 mm., and a spire angle of 83°; another specimen is almost 33 mm. long. Some characters are present that are different from the form found in the Paris Basin, not enough to make it a distinct species, but sufficient to con- sider it a well-defined variety. The principal dif- ferences are the following: the form is much more oblong than the figure given by the worthy De- shayes [this is H. mutica var. a Deshayes, possi- bly the same as E. raricostata (Risso, 1826)], but more like that given by Lamarck (p. 40, pl. 44. Mem. sur les foss. des environs de Paris); ribs of the body whorl usually ten or eleven; between them an elegant, net-like, cancellate sculpture, which is formed by numerous spiral threads and fewer (about 5), more prominent axial ones that resemble sometimes little riblets. Neither De- shayes nor Lamarck note this difference, but the figure of the latter author distinctly shows the axials while the spirals are lacking; evidence of the supremacy of the former over the latter. The number of whorls is six or seven; the first smooth, Plate 194. Eocithara mutica hilarionis Gregorio, 1880. Middle Eocene of Italy. Holotype. 35 mm. in length (from Gregorio, 1880, pl. 5, fizs. 43a, b). submamillate; the last whorl large, but not as globose as that figured in the work of Deshayes in which the posterior convexity seems to cover part of the spire. This species seems to have been found at Croce Grande by Bayan, and listed in his Cata- logue as Harpa cf. mutica Lamarck.” Synonymy— 1880 Harpa mutica var. Hilarionis Gregorio, Fauna di S. Gio- vanni [larione (Parisiano). Parte 1. Cefalopodi e Gas- teropodi, p. 42, pl. 5, figs. 43a, b. Eocithara clarki (Vokes, 1937) (PI. 195) Range—Middle Eocene of California (Domen- gine state). Remarks—This species is more narrow than any other known species of Eocithara, with a higher spire, and without any secondary spiral sculpture. Description (copied from Vokes, 1937)—Holo- type: Univ. California Mus. Paleont. 15792; loc. A-1165, Big Tar Canyon, south of Coalinga, Cali- fornia; Domengine formation. Dimensions: Length, 26.4 mm.; diameter, 14.6 mm. Shell of moderate size, thin, subfusiform; spire high, of five post-nuclear whorls, ornamented with slightly sinuous, lamellar axial ribs separated by broad interspaces marked by five narrow, second- ary axial threads and numerous spiral ribs of equal strength, separated by interspaces about twice the width of the spiral; with 13 axial lamel- lae on the third post-nuclear whorl, 14 on the penultimate, and 16 on the ultimate whorl; whorls shouldered but slightly, sutured appressed, dis- tinct; aperture twice the length of the spire, nar- Plate 195. Eocithara clarki (\iokes, 1937). Middle Eocene of California. 26.4 mm. (from Vokes, 1937, pl. 2, figs. 5, 7). [20-626] November 27, 1973 row; outer lip simple, inner lip with a characteris- tic callus wash; base of the shell concave, recurved to a strong fasciole ornamented with the axial lamellae. Synonymy— 1937 Harpa (Eocithara) clarki Vokes, Jour. Paleont., vol. 11, p. L1, pl. 2, figs. 5, 7. Eocithara raricostata (Risso, 1826) Range—Upper or Middle Eocene of southern France (Alpes Maritimes). Remarks—This species, very briefly described and unfigured, has been ignored by all later workers. Judging from the short diagnosis, it is characterized by its broad inflated shape and dis- tant axial ribs. It may be that this is the same as the Harpa mutica var. a of Deshayes (1824, Descr. Coq. Foss. Env. Paris, vol. 2, p. 642, pl. 86, figs. 14, 15) which has the same characteristics, and was found at Parnes in the Paris Basin. Only an examination of the type, if it exists, can determine its relationship with the other known Eocene species. Synonymy— 1826 Harpa raricostata Risso, Hist. Nat. Europe Merid., vol. 4, p. 180 (“Calcaire grossier de nos montagnes’ ). Eocithara elegans (Deshayes, 1835) (PI. 196) Range—Upper Eocene of France (Auversian of Paris and Nantes Basins). Remarks—This species is somewhat less inflated than E. mutica (Lam.), and is described as posses- sing a sharper subsutural angle on the body whorl. The figures, however, given by Cossmann and Pissaro (1913, pl. 46, figs. 209-2) show a shell Plate 196. Eocithara elegans (Deshayes, 1835). Upper Eocene of France. X 1. (from Deshayes, 1835, pl. 86, figs. 16, 17. INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Eocithara = 227 with a rather rounded subsutural shoulder. The spiral sculpture is stronger than in E. mutica. E. elegans shows some relationship with E. sub- mutica Orb. Description (freely translated from original diagnosis)—This Harpa is ovate-oblong, a little more cylindrical than the preceding species [H. mutica Lam.]; its spire is short: with six very narrow [= low] whorls of which the last are flat- tened above. There are thirteen or fourteen longi- tudinal ribs on the last whorl; these ribs are thicker than in the preceding species, and they are elon- gated a little in their upper part, as in Harpa nobilis. The intervals between the ribs show rather strong distant, unequal transverse striae, a finer one being situated between the stronger ones. These striae form an elegant network with other regular and much finer longitudinal ones. The aperture is proportionately larger than in the other species; the left border is much more nar- row, especially at the base of the columella where it leaves uncovered the oblique and scaly fasciole which ends at the terminal notch of the aperture. The largest specimen that we know of this rare and precious species is 33 mm. long and 20 mm. wide. Synonymy— 1835 Harpa elegans Deshayes, Descr. Coq. Foss. Env. Paris, vol. 2, p. 643, pl. $6, figs. 16-15; 1844, Deshayes, Hist. Nat. An. s. Vert., Ed. 2, vol, 10, p. 134 (Valmon- dois, France). 1913 Harpa (Eocithara) elegans, Cossmann and Pissarro, Icon. Compl. Coq. Foss. Eocene Eny. Paris, vol. 2, pl. 46, fig. 209-2. Eocithara jacksonensis (Harris, 1896) (PI. 197) Range—Upper Eocene of Mississippi (Jackson formation). Plate 197. Eocithara jacksonensis (Harris, 1896). Upper Eocene of Mississippi. 29.5 mm. (from Harris and Palmer, 1947, pl. 56, figs. 19, 20) {20-627 | 228 Eocithara H. A. Rehder Harpidae Remarks—This species is a typical member of the genus Eocithara. The ribs are fewer in number and heavier than in E. mutica (Lamarck), with the intercostal sculpture more pronounced. I have seen two rather poor specimens of this species in the collection of the National Museum of Natural History, of which one measures 31 mm. in length and 18 mm. in width. The holotype measures: length, 31.5 mm.; width 18.7 mm. Description (copied from Harris, 1896)—“Size and general form as indicated by the figure; volu- tions 8; 1 and 2 very minute, smooth; 3 much larger, smooth; 4 somewhat larger than 3, show- ing vertical costae in its first half, then assuming the characteristic markings of the remaining whorls; costae on the body-whorl nine in number, somewhat deflected below the suture as in Drillia; between the costae the shell is finely cancellated with a network of raised lines; anterior canal slightly larger than usual for the genus. Locality, Jackson, Miss.” Synonymy— 1896 Harpa jacksonensis Harris, Proc. Acad. Nat. Sci. Phila- delphia, vol. 48, p. 472, pl. 15, fig. 10. 1947 Harpa (Eocithara) jacksonensis Warris, Harris and Palmer, Bull. American Paleont., vol. 30, no. 117, p. 397, pl. 56, figs. 19, 20. Eocithara birmanica (Vredenburg, 1923) (PI. 198) Range— Upper Eocene of Burma (Yaw stage). Remarks—This species, very briefly described by the author, is based on an incomplete specimen. A plaster cast of the holotype, kindly sent to the National Museum of Natural History by M. V. A. Sastry of the Geological Survey of India, measures 19.1 mm. in length and 12.0 mm. in width. The type was collected at Thetkegyin, Burma. Plate 198. Eocithara birmanica (Vredenburg, 1923). Upper Eocene of Burma. 19.1 mm. left-hand figure, plastocast of holotype; right-hand figure from Vredenburg, 1923, pl. 14, fig. 6 Description (copied from Vredenburg, 1923)— “Although the solitary available specimen is very incomplete, its distinct characteristics make it worthy of record. It is very closely related to Harpa mutica Lamk. of the Middle Eocene of the Paris region, from which it is distinguished by its smaller size and the much more delicate intercostal reticu- lation. The latter character distinguishes it. still more decidedly from Harpa Morgani C. and P. oc- curring in the Lower Eocene of Sind. Compared with Harpa narica Vred. from the Oligocene of Sind, the Burmese shell is smaller, with a rela- tively taller spire and with wider-spaced axial lamellae.” Synonymy— 1923 Harpa (Eocithara) birmanica Vredenburg, Rec. Geol. Survey India, vol. 54, p. 252, pl. 14, fig. 6 (Thetkegyin, Burma). Eocithara submutica (Orbigny, 1852) (PI. 199) Range—Lower Oligocene of southern France (Dax, Landes). Remarks—I have been unable to find a descrip- tion of this species which is based on a name ac- companying a pair of figures. These original figures show a great resemblance to the figures for E. elegans (Deshayes) from the Upper Eocene of the Paris Basin. I have found no references to this species later than that by Cossmann (1899, p. 76) who assigns it to the Tongrian (lowermost Oligocene of Upper Eocene); some later workers place the beds whence this species is supposed to come in the Stampian, or Middle Oligocene. Synonymy— 1845 Harpa mutica Lamarck, Grateloup, Conch. Foss. Tert. Bassin Adour. vol. 1, Atlas, 1840-1845, suppl. pl. 1, fiys. 21, 22 (Dax, Landes, France). Plate 199. Eocithara submutica (Orbigny, 1852). Lower Oli- gocene of France. X 2. (from Grateloup, 1845, pl. 1, figs. 21, 22). [20-628] November 27, 1973 1852 Harpa submutica Orbigny, Prodr. Paléont., vol. 3, p. 17. New name for H. mutica Grat., 1845, not Lamarck, 1803. 1899 Harpa (Eocithara) submutica Orbigny, Cossmann, Es- sais Paléoconch. Comp., pt. 3, p. 76. Eocithara narica (Vredenburg, 1925) (Pl. 200) Range—Oligocene of Pakistan (Nari formation). Remarks—The figures given by Vredenburg are poor, but I have been able to examine plaster casts of the syntypes of this species. These together with his lengthy description suggest that narica has some resemblance to certain species of Harpa, especially in the ovate shape of the body whorl without a shoulder angulation, and in the apparent junction of the forward curving upper ends of the successive axial ribs. Indeed, Vredenburg suggests that this species may represent an ancestral form of Harpa conoidalis (= major Riding). Neverthe- less, I am retaining this species in Eocithara be- cause of the restricted and marginate parietal and columellar callus, the uncertainty over the nature of the protoconch, and because of the age of the beds in which the species is found. Description (abbreviated from the original de- scription)—The elongate-ovoid shell has a low, broadly conical spire, the body whorl measuring nine-tenths of the total length. The small eroded protoconch is followed by three and a half convex whorls, the sutures of which are covered by the junction of the forward curving upper ends of the successive axial ribs. These number from 15 to 16 on the spire whorls, and from 14 to 15 on the body whorl, and show a slight spinosity at the rounded shoulder of the whorls. Between the ribs is a deli- cate network of fine axial and spiral threads. The parietal callus is conspicuously margined by a Plate 200. Eocithara narica (Vredenburg, 1925) Oligocene of Pakistan. 29.0 mm. Plastocast of syntype, USNM. [20-629] INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Eocithara — 229 low, raised edge, and the rectilinear edge of the columellar callus is slightly raised at the pseud- umbilical chink on the inner side of the siphonal fasciole. Measurements (in mm.)— Height 29.3 41.0 Diameter 17.0 24.0 Height of spire 5.4 6.5 Height of body whorl 26.5 38.0 Synonymy— 1925 Harpa (Eocithara) narica Vredenburg, Mem. Geol. Sur- vey India, vol. 50, p. 122, pl. 1, fig. 16, pl. 2, fig. 6 (Bhagothoro Hill, Sind, Pakistan). Eocithara bellardii (Sacco, 1890) (PI. 201) Range—Oligocene of northern Italy. Remarks—Although the brief diagnosis (copied in free translation below) does not mention any characters that assist us in determining whether this species is an Eocithara or a Harpa, the illus- tration given by Sacco does show the narrow pari- etal and columellar callus, the strongly angulate body whorl, and ribs that are apparently not strongly curved forward at the sutural margin. For these reasons I am placing the species in the genus Eocithara. Description (from Sacco, 1890)—Shell of med- ium size, oblong ovate. Spire fairly high. Whorls about six, with heavy, somewhat rounded, very elevated longitudinal ribs; 12-13 subarcuate ribs in last whorl, which generally alternate with the ribs of the penultimate whorl. Surface between the ribs sculptured with numerous very fine trans- verse striae. Aperture subfusiform. Outer lip slightly thickened. Columella subumbilicate. Si- phon somewhat elongate. Height 30 mm., width 18 mm. Synonymy— 1890 Harpa bellardii Sacco, Moll. Terr. Terz. Piemonte Li- guria, pt. 7, p. 9, pl. 1, figs. la, 1b (Tongrian of Cas- sinelle, Dego, Mornese). Plate 201. Eocithara bellardii (Sacco, 1890). Oligocene of north- ern Italy. 30 mm. (from Sacco, 1890, pl. 1, figs. la, Ib). 230 Eocithara H. A. Rehder Harpidae Eocithara muticaeformis (Martin, 191€) (Pl. 202) Range—Lower Miocene of Java (Gunung Spo- long, West Progo Mountains. Remarks—This species appears to be the latest representative of Eocithara, with which it agrees, as Martin points out, in the nature of the parietal callus, the umbilical chink, and the character of the junction of the ribs at the suture. It is begin- ning to show some relationship with species of Harpa in that the outer edge of the parietal callus is thin and is not sharply margined at the outer edge. Description (freely translated from original)— Shell ovate, somewhat inflated, anteriorly attenu- ate. The protoconch button-shaped, consisting of two small smooth whorls, the intermediate whorls strongly convex, with high, narrow, axially ori- ented, slightly curved ribs which in the early whorls develop a bluntly angulate point near the suture. That makes it appear as if a spirally ori- ented angulation, actually absent, were present {on the body whorl]. Between the distant ribs the surface is covered with fine spiral ridges; they are also separated and number up to six. The body whorl has fourteen ribs which are curved forward at the suture which they almost cover; a spiral sculpture similar to that on the spire whorls, is present only on the sutural ramp. [20-630] The columella is slightly concave in the middle; the inner lip very thin but bordered in its anterior half by a definite furrow, and with the margin distinctly erect over the weakly indicated umbili- cal chink. The latter is surrounded by a strong spiral fasciole conforming to the siphonal canal, and over which the axial ribs continue in undimin- ished strength in a sickle-like curve. The outer lip is missing. Length 20 mm. Synonymy— 1916 Harpa (Eocithara) muticaeformis Martin, Samml. Geol. Reichs-Museums Leiden, N.F., vol. 2, pt. 6, p. 231, pl. 1, figs. 15, 15a (Gunung Spolong, West Progo Mts., Java). Plate 202. Eocithara muticaeformis (Martin, 1916). Lower Miocene of Java. 20 mm. (from Martin, 1916, pl. 1, figs. 15, 15a). November 27, 1973 Eocithara species (PI. 203) Range—Lower Eocene (Mangaorapan stage) of New Zealand. Remarks—This specimen is of interest as it rep- resents the second oldest known species of the Harpidae, and also because it shows the wide range that Eocithara had in Lower and Middle Eocene times, throughout the extent of the Tethys Sea, from California, across Europe and India, to New Zealand. Although new species have been described from poorer specimens than this one, it is inad- visable to give this single specimen a name. I am merely putting it on record to fill out the distribu- tional picture of the genus. The figures adequately show that the characters of the shape of shell and of the ribs appear to be those of a typical Eoci- thara. Shell length 27.9 mm., width 15.2 mm. Locality—Tuffs in bed of Whit’s Creek, Eyre River District, Canterbury, New Zealand. Collected by J. Gellen, January, 1960. Age—Mangaorapan stage—equivalent of about upper Ypresian stage; Lower Eocene. The above information and the photographs were kindly sent me by Dr. Alan Beu of the New Zealand Geological Survey. Plate 203. Eocithara species. White’s Creek, Eyre River Dis- trict, Canterbury, New Zealand. Lower Eocene (Mangaorapan stage). New Zealand Geological Survey Eocithara species (Pl. 204) Range—Middle Eocene of Texas (Claiborne for- mation of easternmost Texas). Remarks—The fragmentary specimen is pre- sented here to indicate the presence of a species of INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Eocithara = 231 Plate 204. Eocithara sp. Middle Eocene of Texas. 14 mm. (from Palmer, 1937, pl. 65, fig. 6). Eocithara in the Middle Eocene of North America. It has wider ribs and somewhat weaker intercostal sculpture than does E. jacksonensis (Harris) from the Jackson formation of the Upper Eocene. The locality is in eastern Sabine Co., Texas, on the edge of the Sabine River, east of Hemphill. Synonymy— 1937 Harpa sp. Palmer, Bull. American Paleontology, vol. 7, no. 32, p. 398, pl. 65, fig. 6 (eastern Sabine Co., Texas). Eocithara species Range—Upper Miocene of Eniwetok, Marshall Islands. Remarks—A fragment 15 mm. long from a drill- hole (No. E-1) on Parry Island, Eniwetok, Marshall Islands, is referred with some doubt to this genus. Only the anterior canal and sinus, the lower por- tion of the columellar lip, and a part of the body whorl comprising three complete ribs from their procurved upper end to the siphonal fasciole and anterior canal is present. My assignation of this fragment to Eocithara is based on the low, not broadly expanded, procurved end of the ribs, and the nature of the siphonal fas- ciole separated from the lower end of the columellar callus by a deep furrow which suggests a pseud- umbilical chink. The fairly crowded ribs are low, somewhat flat- tened in their upper portion, with faint, paired color lines visible under magnification. The inter- spaces show flattened spiral cords crossed by irregular axial growth striae. The crowded micro- scopic axial striae present in Harpa major Roding are absent, and the aspect of the sculpture is gen- erally distinct from that found in the recent species of Harpa. This fragment was found in the portion of a drillhole recovered from 830-840 feet, and is as- signed by Ladd (1966, p. 7) to the Upper Miocene. Further, more complete material is needed to prove the generic allocation. If this placement is correct it is the latest known occurrence for Eoci- thara. strongly convex on the ventral surface and [20-635 ] 232 Eocithara H. A. Rehder Harpidae a a a ee [These occasional blank areas occur be- tween genera and subgenera to permit the insertion of new material and future sections in their proper systematic se- quence. | [20-636] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Eocithara = 233 Subgenus Marwickara Laws, 1935 Type: Eocithara (Marwickara) waihaoensis Laws, 1935 This subgenus was proposed by Laws for a single species from the Middle Eocene of New Zealand. It had been sent to Dr. J. Marwick for examination whose summary of the differences between the New Zealand species and the type species of Eocithara are quoted by Laws. The principal differences distinguishing — this subgenus are the more narrowly ovate shape and higher spire, a protoconch with more flattened, less convex whorls, and a neck and anterior canal that is somewhat twisted to the left. The parietal “denticle” seen on the parietal wall about a third of the distance down from the posterior angle with the outer lip, with a lower obscure swelling below it, may be due merely to an irregularity covered over by the parietal wall. Synonymy— 1935 Marwickara Laws, Trans. Royal Soc. New Zealand, vol. 65, p. 28 (Type by original designation: Marwickara waihaoensis Laws). Eocithara waihaoensis Laws, 1935 (Pl. 205) Range—Middle Eocene of New Zealand. Remarks—The following description is based on the holotype and two fragments kindly sent me on loan by Mr. Walter O. Cernohorsky of the Auck- land (New Zealand) Institute and Museum. Description—Shell relatively small (23 mm. in length), ovate, with rather high spire (more than one third of total length, as compared with less than one fourth of total length in Eocithara). Protoconch somewhat worn but showing rather flattened nuclear whorls. Early — post-nuclear whorls also somewhat worn, with fairly strong opisthocline ribs; maximum curvature at shoulder of whorls; ribs numbering twenty in number in penultimate whorl, bluntly angulate in cross-sec- tion, not lamellar, with interspaces marked by strong, subequal spiral cords crossed by irregular growth wrinkles; the anterior ends of the ribs form only low ridges on the weakly differentiated si- phonal fasciole. Aperture rather narrow, outer lip thickened internally, somewhat reflected, sinuous in profile with a shallow posterior sinus and proso- cyrt medially. Inner lip with a distinct and slightly thickened outer margin, which in the type is bro- ken. Siphonal canal moderately long, directed to the left. Specimens examined—Holotype (Auckland Inst. and Museum): length, 23 mm., width 12.3.mm.; greensands, Waihao Downs, South Canterbury, New Zealand (Bortonian, Middle Eocene). Para- types: two fragments from same collection. Synonymy— 1935 Eocithara (Marwickara) waihaoensis Laws, Trans. Royal Soc. New Zealand, vol. 65, p. 29, fig. 11 (Waihao Downs, South Canterbury, New Zealand). Plate 205. Eocithara (Marwickara) waihaoensis Laws, 1935. Holotype, Waihao Downs, South Canterbury, New Zealand; [20-641] Middle Eocene. 23 mm. Auckland Institute and Museum. 234. Marwickara H. A. Rehder Harpidae [These occasional blank areas occur be- tween genera and subgenera to permit the insertion of new material and future sections in their proper systematic se- quence. | [20-642] November 27, 1973 Subgenus Refluharpa Iredale, 1931 Type: Harpa lamellifera Tate, 1559 This subgenus contains only one species, E. (Refluharpa) lamellifera Tate from the Middle Miocene of Victoria. Iredale proposed the genus Refluharpa for this species, which Finlay in the same year had placed in Eocithara. Cotton and Woods (1933, pp. 45, 47) made Refluharpa a synonym of Eocithara because of the similarity of the protoconch. However, the protoconch of E. lamellifera is considerably larger than that of E. mutica La- marck; the maximum diameter of the protoconch of a specimen of the former being more than twice as great as that of an example of E. mutica; the number of whorls is generally greater 2% to 3 as opposed to 2% in mutica; the apex is more pla- nate with the suture more deeply impressed; the parietal callus is not strongly marginate but is thin and evanescent near its outer edge. These differences plus the later age of E. lamel- lifera, induce me to retain the species in a distinct subgenus. Synonymy— 1931 Refluharpa Iredale, Rec. Austral. Museum, vol. 15, pp. 230, 233 (June 29). Eocithara lamellifera (Tate, 1889) (Pls. 206, 207) Range—Middle Miocene (Balcombian-Bairnsdal- ian) of Victoria. Plate 206. Eocithara (Refluharpa) lamellifera (Tate). Proto- conch of specimen from Muddy Creek, Hamilton, Victoria; Middle Eocene. USNM 646909. X 10. INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 bo oe ur Eocithara Remarks—In addition to the differences listed above under the subgenus, this species is marked by the great number of strongly lamellar ribs— more than 35 on the body whorl of a specimen 24.5 mm. in length, by the subsuturally planate whorls, the prominent spiral sculpture between the ribs which is particularly prominent in the penultimate whorl. It is found in several localities in Victoria— Mud- dy Creek, Hamilton (the type locality); Balcombe Bay, Port Phillip Bay; near Altona, Port Phillip Bay; Shelford, near mouth of Gellibrand River. Measurements (mm. )— length width 32 21.5 Hol type (SAM-T-698) 36 22 Paratype (SAM-T-698) 24.5 16.1 USNAIL 646909 23.6 15.9 USNM Synonymy— ISS9 Harpa lamellifera Tate, Trans. Proc. Rep. Royal Soc South Australia, vol. 11, p. 149, pl. 6, fig. 2 (lower beds, Muddy Creek, Victoria) 1897 Harpa (Eocithara) lamellifera Vate, Harris, Cat. Tertiary Moll, Dept. Geol. British Museum, pt. 1, p. 79, pl. 4, fig. 3 .a-b. 1931 Eocithara lamellifera Tate, Finlay, Trans. New Zealand Inst., vol. 62, pp. 12, 13 (May 31); 1933, Cotton and Woods, Records South Australian Museum, vol. 5, pp. 45, 47 1931 Refluharpa lamellifera Yate, Iredale, Records Australian Museum, vol. 18, p. 230 (June 29), Plate 207. Eocithara (Refluharpa) lamellifera (Tate). Holo- type (30.5 mm.) and paratypes. South Australian Museum, Tate Colln. 698. [20-647] 236 Refluharpa H. A. Rehder Harpidae [These occasional blank areas occur be- tween genera and subgenera to permit the insertion of new material and future sections in their proper systematic se- quence. | [20-645] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpa 237 Genus Harpa Roding, 1798 Type: Harpa harpa Linné, 1758 The genus Harpa comprises nine recent and five fossil species. Seven of the living species are found in the Indo-Pacific region and one each on the West African coast and in the eastern Pacific. The fossil species are found in beds of from Oligo- cene to Pliocene age. The shells are characterized by their relatively large body whorl bearing a variable number of axial lamellar or sublamellar ribs, the rather ex- tensive parietal and columellar callus which is not sharply marginate abaperturally, and by the whorls of the spire, especially the penultimate and antepenultimate whorls, being more or less covered by the glaze of the expanded posterior part of the axial ribs. The protoconch is elevated- conic, consisting of from 3 to 5 whorls, usually flesh-pink to purplish red, sometimes whitish in color, with a distinct keel at the periphery, just visible above the suture. The basic color pattern is a banded one, the axial ribs marked with al- ternating spots of various shades of pink and white, with or without dark horizontal lines, these spots coinciding on successive ribs; the banded effect thus created is strengthened by the irregular, axial, zigzag and festooned dark lines and white spots in the spaces between the ribs, this pattern also being repeated in the suc- cessive interspaces, with the adapertural projec- tions of the festoons always coinciding with the white spaces on the axial ribs. In Harpa costata much of the color pattern is obscured by the dense axial sculpture. The anterior siphonal notch is rather broad and open, not narrow and somewhat constricted as in Eocithara. Synonymy— 1798 Harpa Roding, Museum Boltenianum, p. 149 (type by tautonymy: Harpa harpa Linne). 1799 Harpa Lamarck, Mem. Soc. Hist. Nat. Paris, vol. 1, p. 71 (type by tautonymy: Harpa harpa Linne). 1806 Harpalis Link, Beschr. Nat.-Sammlung Univ. Rostock, pt. 3, p. 114 (type here designated: Harpa major Lamarck). 1815 Harparia Rafinesque, Analyse de la Nature, p. 145. New name for Harpa Lamarck, 1799. 1934 Lyra Griffith and Pidgeon, Cuvier’s Animal Kingdom, vol. 12, p. 234. Nomen nudum. 1881 Cithara “Klein” Jousseaume, Bull. Soc. Zool. France, vol. 5: Proc.-Verb., p. xxxviii (type here designated: Harpa harpa Linne). Nomenclature—Some authors have credited the name Harpa to Walch, 1771, or have cited an earlier usage of the name, namely Harpa Pallas, 1774. The first is based on Herrmannsen’s cita- tion (Herrmannsen, 1846-47, p. 501), who gives, however, an erroneous reference. On page 113 of volume 2, part 1 (not volume 3, p. 113), Walch discusses the “Harfenschnecken” but without us- ing a Latin name; furthermore, Walch’s work is non-binominal. Pallas in his “Spicilegia Zoologia” used Harpa nobilis as a vernacular group name in connec- tion with his description of Buccinum geversi- anum (=Trophon g.). As such, in addition to being in the plural form, it is not available as a validly proposed taxon. Harpa harpa (Linne, 1758) (PI. 187, figs. 7-10) Range—From East Africa to Tonga. Remarks—This species is characterized by its stout, broadly ovate, markedly shouldered shell, with three separated chestnut blotches on the ventral side, the ribs rather strong, flattened, and marked with many distinct lines arranged in groups; a band of interrupted and_ irregular blotches of orange brown or reddish brown is generally present about the middle of the body whorl. Plate 208. Enlarged protoconchs of species of Harpa. Fig. 1. Harpa ventricosa Lamarck. USNM 742la. Fig. 2. Harpa arti- cularis Lamarck. Off Tambizen, North Borneo, USNM 666808. Fig. 2. Harpa harpa (Linné). USNM 7421. Fig. 4. Harpa cre- nata Swainson. Mulege Bay, Baja California, Mexico, USNM 12509. (all X 5). | 20-653 | 238 Harpa H. A. Rehder Harpidae 60° 80° 100° 120° 140° 160° 180° 160° — T 7 a3) ] T T + 7 eee | | | a || Hochyjo a | | $ 2 i | I >: i Midway,» Pearl f Hermes Nt , P ¢ BONIN f pair ah| oflono > ers MAwa, | , -€ Bie Z 1a ~—— ES -|- Re se a NE a a cre en ae SS ites ee eae Is | FORMOSA ae ly | , : Oahu Os —— =e) = Suman ies = Be = ae a] — {ES | i MARIANA Weke Hawai Ms - Johnston ' : PHILIPPINE eg : = . \PP\ , | * 7 Z @ ht 4 2p % is a ink, Enwetohk@ - iD RSHALL IS NICOBAR; t yt Polou Y eer er, ae CR tad : E 5° i, A Meet a a) Me, anaes. “+ Prajuro pe % cEYY Saar CAROLINE is! [o> ee pails MALDIVE » ~ ti a | ° : | F s Mak <.. |" EDIE 2 . | ‘Helen Rf cm ial 4 | = a 2 ka 4 | ©. | ares eo 9 a = 7 AVY 3. fe SAYCHELLES 1S. » oy nt = ‘e XI Sy : ’ |e io CHAGOs |” ae * (PHOENIX 15 ig Gas : ye UA “ems . TOKELAU 1S ert | OCG - KEELING 5 2 L 3 ‘ss . | ve | ATOLLS “ | SA, v . p= ' A a we New “@ 3 eI ea. G PHEBRIDES ud | N > 6 4 wv Le S| J j Qo onew \.a¥itino M3 1s. . 20° 20° 60° Plate 220. Geographical distribution of Harpa_ ventricosa Lamarck. H. A. Rehder Harpidae the parietal wall. A third small spot is present at the base of the columellar lip and is occasionally connected to the lower large spot by a chestnut patch along the inner edge of the columellar lip. Aperture ovate, outer lip gently rounded or occa- sionally somewhat flattened; interior usually with yellow-orange coloration, and with the external banded pattern visible. Measurements (mm.)— length width no. whorls 108.8 78.3 large; Mauritius 99.5 67.7 7% large; Seychelles 85.3 58.4 8 average; Mauritius 55.3 38.7 64 small; Zanzibar 47.8 32.6 small; Zanzibar Synonymy— 1816 Harpa ventricosa Lamarck, Encyclopédie Méthodique, vol. 3, pl. 404, figs. la, 1b., Liste, p. 3; 1822, Lamarck, Hist. Nat. Anim. sans Vert., vol. 7, p. 255 (Mers des Indes orientales); 1835, Kiener, Coquilles Vivantes, vol. 8, Gen. Harpe, p. 6 (in part), pl. 1, fig. 1, pl. 4, fig. 7; 1843, Reeve, Conchologia Iconica, vol. 1, Harpa, pl. 1, sp. 2 (in part: figs. 2b, 2c, 2d only); 1857, Kuster, Neues Syst. Conch.-Cabinet, ed. 2, vol. 3, pt. 1, p. 89, pl. 67, figs. 1-3; 1860, Sowerby, Thesaurus Conchologia, vol. 3, p. 169 (in part), pl. 232, figs. 18- 22, pl. 233, fig. 25; 1877, Sutor, Jahrb. deutsch Malak. Ges., vol. 4, p. 99. 1822 Buccinum harpa var testudo Donovan, Naturalist’s Re- pository, Exotic Natural History, vol. 1, pl. 8. Not B. testudo Lightfoot, 1786, a nomen dubium. 1843 Harpa conoidalis Lamarck, Reeve, Conchologia Iconica, vol. 1, Harpa, pl. 3, sp. 7 (in part): fig. 7b only; not Harpa conoidalis Lamarck, 1822). 1860 Harpa cabritii Fischer, Journ. de Conch. vol. 8, p. 209, pl. 4, figs. 1, 2 (juvenile) (no locality). 1948 Harpa major Réding, M. Smith, Triton, Helmet and Harp Shells, p. 48 (in part), pl. 16, fig. 7 (not Harpa major Réding, 1798). Types—In the Museum d Histoire Naturelle in Geneva no specimens were found in the Lamarck Collection that agree exactly with the figure in the Encyclopédie Méthodique. In the Delessert Collection, however, I found a specimen that agrees with the above-mentioned figure but is slightly smaller (96 mm. in height instead of 100.4). This specimen I designate as the neotype. The type of H. cabritii Fischer is in the British Museum (Natural History), catalogue number B. M. (N. H.) 99.8.22.126. The whereabouts of the type of H. testudo Donovan is not known. Records—SOUTH AFRICA: off Durban, Natal. from fish (Colln. Helen Boswell). MOZAMBIQUE: Mogambique (ANSP, BM, CMNH); Porto Amelia (AMNH, DMNH, MCZ). TAN- ZANIA: Dar es Salaam (MNH); Mijimwenda, 5 mi. ESE of Dar es Salaam (MCZ); Zanzibar (ANSP, BM, DMNH, MCZ, RNHL); Kiwengwa, in 0 to 10 ft.; Chumbe Id., 0 to 6 ft.; Pange Id., 0 to 2% ft.; Mnemba Id.; 2 mi. W of Bani Id., 15 fms. (all Zanzibar and ANSP). KENYA: Mombasa (BM); Diani Beach; Lamu Id. (both MCZ); Tiwi (MHNG): RED SEA Straits of [20-668] November 27, 1973 Jubal (ANSP). ADEN (BM). SOCOTRA: north coast (ANSP). MADAGASCAR: between Ambotoloaka and Madioranokely, SW of Nosy Bé; Amforah, Nosy Be (both ANSP); S of Ambo- vombé (AMNH); Nosy Faly (FNHL). ILE GLORIEUSE: (USNM). SEYCHELLES: (MCZ); Mahe (BM). MAURITIUS: (AMNH, ANSP, BM, DMNH, MCZ, USNM); Mahebourg (USNM); Ile Flamand (DMNH). Doubtful Records—Tranquebar Coast, India (RNHL); Ma- dura (RNHL) and Ambon, Indonesia (MCZ, RNHL); these three records are based on specimens from old collections, and should be regarded as doubtful. No specimens of this species have been found in India or Indonesia in recent years. Two specimens in the USNM are labeled as coming from Luzon and from Negros Oriental; the localities of these speci- mens, obtained from a dealer and from an amateur collector respectively, are suspect. Seat ees Harpa costata (Linne, 1758) (Pl. 187, figs. 1-3; Pls. 221-223) Range—Islands of the western Indian Ocean: Mauritius, Rodrigues, and northeastern Mada- gascar. Remarks—This is one of the most distinctive species, characterized by its broad shape, the large number of crowded ribs, which are rather sharply pointed at the shoulder-angle of the body whorl forming a broad subsutural channel; its dis- tinctive coloration consists of numerous. spiral bands of varying shades of flesh color and white. Habitat—On sand banks, frequently in shallow water—10 inches to 6 feet. Description—Shell 70 to 100 mm. (2% to 4 inches) in length, broadly subquadrate, solid, last whorl very large. Spire broadly conical; protoconch ele- vated-conical, flesh pink, whorls 44, smooth; early postnuclear whorls broadly shouldered, with spaced axial riblets and fine spiral cords below the shoul- der and light chestnut spots on the shoulder be- tween the protractively curved riblets. On the later postnuclear whorls the axial riblets become more crowded, and on the shoulder become increasingly lamellar towards the upper portion where they are adnate to the preceding whorls, fusing and cover- ing the lower half of the exposed whorls; in the last half of the penultimate whorl the spiral cords become obscure, in some specimens completely covered by the fused upper part of the ribs of the following whorl. On the body whorl the ribs are crowded, shiny, 30 to 40 and more in number, de- pending on the size of the shell; they are lamellar, recurved, with a triangular, spinelike projection at the shoulder angle, forming a broad, subsutural channel; often more or less flattened in the last half of the body whorl, and fused above the con- spicuous siphonal fasciole over which the lamellar INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpa 253 ribs are strongly continued. Between the ribs are fine axial cords crossed by distant, subobscure spiral cords. External color of shell pale flesh color, with bands of varying darker shades and white; occasional subquadrate spots of chestnut color are present between the ribs, usually in an irregular peripheral band. Aperture broad, appear- ing subrectangular because of the broad subsutural shelf; inner lip almost straight, outer lip angled at the shoulder and rather effuse at the base, slightly thickened within; parietal callus rather thin, colu- mellar callus heavier, both usually somewhat suffused with yellow, and with three chestnut blotches, the upper and middle ones moderately large, generally subequal, the lower one on the columellar callus small, obscure or absent; interior of aperture often with a yellow wash, especially near the base. From color slides kindly sent me by Mrs. E. Couacaud of Port Louis, Mauritius, (Plate 221) the foot of H. costata appears to be relatively broader than in either ventricosa or major, and the pos- terior end is rather obtuse with a keel marking the posterior part of the foot. Plate 221. Dorsal view of Harpa costata (Linné) from Mauritius (photo by Mrs. E. Couacaud). [20-669] 254 Harpa Plate 222. Harpa costata (Linné) Holotype of Harpa costata var. laetifica Melvill, 1916. National Museum Wales. 42.1 mm. in length. Measurements (mm.)— length width no. whorls 99.0 78.0 large; Mauritius 97.4 69.4 large; Ile aux Fouquets, Mauritius 92.0 67.2 6% average; Le Morne, Mauritius $5.1 61.0 average; Ile aux Fouquets, Mauritius $1.8 63.9 average; Ile aux Fouquets, Mauritius 71.3 54.8 64 small; Mauritius Synonymy— 1758 Buccinum costatum Linné, Systema Naturae, ed. 10, vol. 1, p. 738 (no locality); type locality here desig- nated: Mauritius; [1788 Harpa imperialis Chemnitz, Conchylien-Cabinet, vol. 10, p. 184, pl. 152, fig. 1452 (no locality); non-binominal.] 1822 Harpa imperialis Lamarck, Hist. Anim. sans Vert., vol. 7, p. 225 (Mers de lAmerique meridionale?); 1853, Chenu, IJlustr. Conchyliologiques, vol. 4, pt. 85, Harpa, pl. 1, fig. 1, la, 1b. 1822 Harpa multicostata Sowerby, Genera of Shells, no. 3, Harpa, fig. 1 (Indian Ocean). 1835 Harpa ventricosa var. Kiener, Coquilles Vivantes, vol. 8, Genre Harpe, »p. 7, pl. 2, fig. 2 (no locality). 1843 Harpa imperialis Chemn., Reeve, Conchologia Iconica, vol. 1, Harpa, pl. 2, fig. 5; 1857, Kuster, Neues Syst. Conchylien-Cabinet, ed. 2, vol. 3, pt. 1B, p. 86, pl. 66, figs. 1-2, pl. 70, fig. 1. 1860 Harpa costata Linné, Sowerby, Thesaurus Conchologica, vol. 3, p. 169, pl. 231, figs. 4-5, pl. 233, fig. 23 (young); 1883, Tryon, Man. of Conch., vol. 5, p. 97, pl. 40, fig. 58. 1877 Harpa costata var. gruneri ‘Maltzan’ Sutor, Jahrb. deutsch. Malak. Gesellschaft, vol. 4, p. 102, pl. 4, fig. 2 (no locality). 1916 Harpa costata var laetifica Melvill, Journ. of Conch., vol. 15, p. 31 (no locality). H. A. Rehder Harpidae Types—There is no specimen of this species in the Linnean Collection in London, and Linnaeus did not cite any reference in his original descrip- tion. According to Odhner (unpublished list and microfilm) a specimen is present in the Museum Ludovicae Ulricae, and this specimen I hereby designate as lectotype. The type locality I am designating as Mauritius. The specimen upon which Chemnitz based his description and figure of his Harpa imperialis and which he stated came “Ex Museo Spengleriano” is present in the Zoo- logical Museum in Copenhagen with a label in Spengler’s handwriting. This specimen I am desig- nating as the lectotype of Lamarck’s species im- perialis; there are no specimens of H. imperialis in the Lamarck collection in Geneva. The type of gruneri Sutor was in the Maltzan collection which according to Dance (1966, p. 293) was purchased by a dealer and dispersed; its present location is unknown. The holotype of Melvill’s variety laeti- fica is in the Melvill-Tomlin Collection in the Na- tional Museum of Wales in Cardiff (Plate 222). Records—MAURITIUS: Le Morne, SW coast (DMNH, USNM); off Ile Marianne and Ile aux Fouquets, W_ coast (ANSP. Colln. W. N. Carpenter); Mahebourg (ANSP, DMNH); Les Benitiers, W coast (AMNH). RODRIGUES (BM). MAD- AGASCAR: 28 km. S of Antalaha, NE coast, in 2 meters (Mme. H. Bouchard, in litt.). 20° a MALDIVE » * 1S. 4 + SEYCHELLES IS. 4) Zonxsbar CHaGee oo ISe 35 comoras . 3 20° 20° e = an | a SFE YET. Plate 223. Geographical distribution of Harpa costata (Linné). [20-670] November 27, 1973 Harpa doris Roding, 1798 (Pl. 189, figs. 12-16; Pl. 224) Range—From the Cape Verde Islands to Luanda, Angola; Ascension Island. Remarks—Harpa doris is most closely related to the only other species found outside of the Indo- Pacific region, namely Harpa crenata Swainson of the Panamic province. This relationship is dem- onstrated by the presence in both species of vivid, spirally oriented, narrow bands of color markings, often in a more or less sagittate pattern, as well as blotches of solid color; in fresh specimens of both species the ribs are marked on their abapertural side by a fine, interrupted chestnut line. Harpa doris, differs from crenata in being smaller, some- what more slender, the ribs with a greater ten- dency to becoming broad, and by the spirally oriented series of blotches being rose or rose- orange rather than chestnut. Description—Shell 77 to 31 mm. (3 to 14 inches) Plate 224. Harpa doris Réding. Holotype of Harpa rosea La- marck, 1816. Muséum d'Histoire Naturelle (Genéve), 55 mm. in length. INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpa = 255 in length, usually elongately oval and rather thin, but occasionally broader and solid. Spire broadly conical; protoconch — elevated-mamillate, 3% whorls, pale pink or flesh-color, smooth; first 1’ postnuclear whorls with low, sharp axial ribs crossing distant rounded spiral ridges; between the ribs the spiral sculpture is crossed by crowded microscopic axial threads. On subsequent whorls the ribs show a spinose angulation at the shoulder, and the spiral sculpture becomes increasingly ob- scure, first on the subsutural ramp above the shoulder, and then on the remainder of the whorl. Body whorl elongate ovate, with a distinct sub- sutural shelf above the shoulder angulation which is marked by the strong triangular spines of the ribs; ribs 11 to 13, occasionally 14, in number, generally slender and rather low, and _ triangular in cross-section; occasionally those towards the aperture are greatly broadened; ribs marked by a thin brown interrupted line on the crest. In the stout broad form, all the ribs on the body whorl are rather broad and heavy and rather angulate at the periphery. Base color pale flesh-pink, occa- sionally darker, with the early postnuclear whorls orange-pink to purplish pink; bands of this color are present on the body whorl below the angula- tion at the shoulder, in the middle of the whorl, and above the base, the latter narrower than the other two; these bands often are bicolorous with alternate squarish blotches of the orange shade and the purplish pink shade; narrow, subequally spaced bands of chestnut marked by sagittate white spots mark the body whorl, these bands oc- casionally linked by obscure, chestnut, arcuate lines. A thin glaze covers the parietal and columel- lar areas which are marked by three separated brown spots: one just above the juncture of the Plate 225. Geographical distribution of: 1, Harpa crenata Swainson, 2, Harpa doris Réding. [20-671] 256 Harpa outer lip with the body whorl, another, the largest, just above where the columella joins the parietal wall, and the third on the expanded columellar lip; the last two spots are separated by the covered siphonal fasciole. Aperture elongate-ovate, semi- lunate, the outer lip gently arcuate. Measurements (mm.)— length width no. whorls 77.2 49.8 6% large; Senegal 49.7 29.0 7% medium; Cape Verde Ids. 48.1 29.5 medium; Cape Verde Ids. 40.1 25.2 small; Annobon 31.1 20.6 small; Fernando Poo Synonymy— [1786 Buccinum pandura ‘Solander’ Lightfoot, Portland Cata- logue, pp. 17, 103 (Guinea). nomen nudum]. 1798 Harpa doris Roéding, Museum Boltenianum, p. 150; refers to Martin, Conchylien-Cabinet, vol. 3, p. 419, pl. 119, fig. 1094 (no locality given; type locality here designated: Accra, Ghana); 1948, M. Smith, Triton Helmet and Harp Shells, p. 47, pl. 16, fig. 4. 1807 Harpalis doris Link, Beschreibung Naturalien Sammlung Univ. Rostock, pt. 3, p. 114. 1816 Harpa rosea Lamarck, Encyclopédie Méthodique, vol. 3, pl. 404, fig. 2; Liste, p. 3; 1822, Lamarck Hist. Nat. Anim. sans Vert., vol. 7, p. 257; 1843, Reeve, Concho- logia Iconica, vol. 1, Harpa, pl. 4, figs. 8a, b, c, d; 1950, Nickles, Mollusques Testacés Marins Cote Occid. d'Afrique, p. 113, fig. 204. Types—Réding based the species doris on a de- scription and figure published by Martini, based in turn on a shell in the latter’s collection. This specimen must be presumed to be lost. The type of H. rosea Lamarck is in the Lamarck Collection in the Museum of Natural History in Geneva. Records—CAPE VERDE ISLANDS: (AMNH, BM, DMNH, MCZ). SENEGAL: (AMNH, BM, DMNH, RNHL). GAMBIA: (AMNH). GHANA: (BM); Accra (AMNH); Takoradi; Elmina (both Buchanan, 1954). SPANISH EQUATORIAL AFRICA: Santa Isabel, Fernando Poo (USNM); Corisco, Rio Muni (AMNH); Annoben (USNM). SAO TOME: (Knudsen, 1956). GABON: Omboue (= Namino, Fernan-Vaz, (Nicklés, 1952): ANGOLA: Santo Antonio de Zaire (Tomlin and Shackelford, 1914); Luanda (MCZ). ASCENSION ISLAND: (DMNH, MCZ); English Harbour (Colln. K. Jourdan); Georgetown Beach, (DMNH). The form generally found washed up on the beaches of Ascension Island is a broader, heavier shell, with broader ribs that are more or less angu- late at the periphery (Pl. 189, figs. 15, 16). I have seen two specimens from Santa Isabel on Fer- nando Poo that also belong to this form. For a time I considered this stout form to be a distinct subspecies restricted to the islands off the African coast. However, I have recently seen a fresh speci- men of the normal form washed up on the sandy beach at English Harbour on Ascension Island. This is one of the few localities on the island where a sandy substrate is present. It is likely therefore H. A. Rehder Harpidae that the normal form occurs where a sand bottom is found, and that in a rocky area where the bot- tom consists of rocks or cobbles, as in most of Ascension Island and on Fernando Poo, the spe- cies develops a heavier shell. The two forms can thus be considered ecophenotypes of the species Harpa doris Roding. Harpa brochoni ‘Benoist’ Cossmann, 1899 (Pl. 226) Range—Late Lower Miocene of France (Burdi- galian). Remarks— This species is a typical Harpa, as evi- denced by the strong development of the parietal callus over the ventral surface of the body whorl and over the lower part of the penultimate whorl. Its closest relative is the West African Harpa rosea Roding, some of whose beachworm specimens closely resemble H. brochoni. The aperture of the latter is shorter, however, with the base less patu- lous, and the knobbing at the shoulder angle is heavier. Description—(translated from Peyrot, 1928)— Shell thick. Size rather large. Form ventricose; spire short, composed of five to six whorls, the first smooth, constituting the protoconch which is badly preserved on my specimens; the following whorls, first convex, then angulate, are orna- mented with a dozen narrow axial ribs, widely separated, subspinose on the angle; from the penultimate whorl they cross the suture and join each other on the preceding whorl; last whorl very large, ventricose, the ribs on it becoming heavy and lamellose; they cross posteriorly the sutural ramp extending onto the preceding whorl while joining each other; anteriorly they curve back- wards hook-shaped over the siphonal fasciole, which thus appears strongly lamellose; the inter- costal spaces show feeble separated spiral striae. Plate 226. Harpa brochoni ‘Benoist’ Cossmann, 1899. Lower Miocene of France. 55 mm. (from Peyrot, 1928 pl. 11, figs. 30-32). [20-672] November 27, 1973 Aperture very dilated, above all anteriorly where it is strongly sinuate; outer lip rectilinear, slightly oblique, with a weak sinus at its junction with the suture, externally thickened by the last rib; inter- nally smooth; columellar margin extensively spread over the ventral surface of the last whorl where it is rather thin, becoming thicker anteriorly where it forms a slight swelling on the edge of the siphonal fasciole, in the columellar area, before terminating in a point on the siphonal notch. Di- mensions: height, 55 mm.; max. diam. 34 mm. Synonymy— [1884 Harpa brochoni Benoist, Proces-Verbaux Soc. Linn. Bor- deaux, 1884, p. LNVII, nom. nud. | P 1899 Harpa brochoni Benoist, Cossmann, Essais, paleoconch. comp., livr. 3, pp. 74, 75, pl. 4, fig. 3 (near Bordeaux, France). P 1928 Harpa brochoni Benoist, Peyrot, Conch. Neogénique Aquitaine, vol. 5, p. 369, pl. 11, figs. 30-32 (Saucats, S of Bordeaux). Harpa josephiniae Sacco, 1890 (Pl. 227) Range—Middle Miocene of northern Italy (Hel- vetian). Remarks—This small species (19 mm. in length) is compared by the author with Harpa ventricosa Lamarck but it is more slender, not as broad as either ventricosa Lamarck or major Réding. In fact it resembles more closely Harpa doris Réding of West Africa, but the ribs are more numerous and are not as spinose below the subsutural ramp, and the body whorl does not show the angulation below the spinose shoulder of that species. The lack of the expanded parietal callus typical of Harpa may be due to the possible juvenile condi- tion of the unique holotype. Description (freely translated from the original) —The following comments distinguish this species from FH. bellardii: Shell smaller, very slightly more ovate. Ribs stouter (especially at the base), less elevated, oc- casionally less numerous; near the suture slightly flattened, above the base generally more widely Plate 227. Harpa josephiniae Sacco, 1890. Miocene of northern Italy. 19 mm. (from Sacco, 1890, pt. 1, figs. 2a, 2b). INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpa 257 separated. Transverse striae nearly obsolete above, below very few, occasionally crossing the ribs. Aperture slightly wider, especially below. Siphon a little broader. Height 19 mm., width 12 mm. Synonymy— 1890 Harpa josephiniae Sacco, Moll. Terr. Terz. Piemonte e Liguria, pt. 7, p. 9, pl. 1, fig. 2a, b (Helvetian of hills near Turin). Harpa americana Pilsbry, 1922 (Pl. 228) Range—Middle Miocene of the Dominican Re- public and southern Vera Cruz, Mexico. Remarks—This species is, as Pilsbry states, close to Harpa doris Réding of the West African fauna, differing in the aperture being narrower anteriorly and in possessing conspicuous fine spiral striation between the ribs; the spiral striation in doris is visible only in certain specimens and even then is rather obscure. H. americana agrees rather closely with H. josephiniae Sacco of the Miocene of north- ern Italy, although the spiral striation is more pronounced in the American species, and the ribs are more spinose at the shoulder. Through the kindness of Dr. Horace C. Richards I have been able to examine the holotype of Harpa americana. As the figure given by Pilsbry agrees in all particulars with the type, and as it clearly shows the essential characters I have reproduced this illustration rather than give a photograph of the type. The characters of the nuclear whorls are not described by Pilsbry nor shown clearly in his fig- ure. The protoconch is erect-mamillate with the first whorl lost; the first 14 of the remaining Plate 228. Harpa americana Pilsby, 1922. Middle Miocene of Dominican Republic. 33.3 mm. (from Pilsby, 1922, pl. 23, fig. 13). [20-673] 258 Harpa whorls shows the basal keel just at the suture that is typical of the protoconch of the genus Harpa. The measurement of the width of the type given by Pilsbry is in error; the true figure is given be- low following the description. Perrilliat (1960, p. 24) describes and figures a slightly larger specimen collected in southern Vera Cruz, 11 kilometers east of Coatzacoalcos; this and another specimen mentioned by her, measure respectively 41 and 37 mm. in height. A smaller shell from the same locality, 26.0 mm. high, is in the collections of the U.S. Geological Survey. Description (copied from Pilsbry)—The shell is ovate, of about 6 whorls, of which three smooth ones form the nipple-shaped embryonic shell, the last whorl of which, together with part of the first sculptured whorl, are very narrow. The last whorl has about eleven low and narrow axial ribs which rise into small spines where they pass over the angle bounding a narrow flattening below the su- ture. The whole surface below this angle is spirally striate, the striation strongest in the concavity of the sides below. The aperture is narrow for this genus. A thin callus spreads forward over the ven- tral convexity. Length 33.3 mm., width 19.4, Dominican Re- public. Holotype, ANSP 4061; length 26.0 mm., width 16.3. mm., Coatzacoalcos-Villa Hermosa Highway, Vera Cruz, Mexico, USGS Colln. Synonymy— 1877 Harpa rosea Lam., Gabb, Trans. American Phil. Society, vol. 15, p. 214. Not H. rosea Lamarck, 1516. 1922 Harpa americana Pilsbry, Proc. Acad. Nat. Sci. Phila- delphia, 1921, p. 337, pl. 23, fig. 13; 1960, Perrilliat, Paleontologia Mexicana No. 8, p. 24, pl. 3, figs. 18, 19. Harpa crenata Swainson, 1822 (PI. 189, figs. 1, 2; Pl. 225) Range—Magdalena Bay, Baja California and southern part of Gulf of California, Mexico, to Gorgona Island, Colombia. Remarks—This species is most closely related to Harpa doris Roding of the West African marine province. These two species are the only living representatives of a small species complex that had its center in the Caribbean area. The rare Harpa americana Pilsbry from the Miocene of the Dominican Republic and Tehuantepec, Mexico is probably close to the ancestral stock of both spe- cies. We may conjecture that this stock once in- habited the West Tethyan Sea, or spread to the H. A. Rehder Harpidae Panamic and West African areas from the Carib- bean, and species became established there while the group died out in the Caribbean. H. crenata differs from doris Roding in being generally larger, broader, the spire relatively lower and broader, the more slender ribs marked more consistently by a fine interrupted chestnut line, and by the squarish blotches being chestnut- brown rather than pink or orange. The body whorl is more markedly angulate be- low the shoulder, the ribs bearing more numerous spines between the subsutural shoulder and the periphery. Habitat—On clay bottom in 40-55 meters (Par- ker, 1964, pp. 155, 172). Description—Shell 32 to 91.5 mm. (1% to 3% inches) in length, broadly oval, body whorl large, more or less strongly angulate at shoulder. Spire broadly conical; protoconch elevated-mamillate, pale corneous, 3% whorls, smooth; axis of proto- conch and first postnuclear whorl sometimes at a slight angle to that of rest of shell; first postnuclear whorl sculptured with distant axial riblets and two spiral cords forming a coarse reticulate pattern which becomes complicated by addition of further spiral cords; in the second postnuclear whorl the upper spiral cord marks an angulate shoulder, the ribs become more lamellar, and the upper termi- nus of the ribs curves forward and forms a layer adnate to lower part of preceding whorl; the ribs at the angulate shoulder are produced into a lamellar, triangular spine; occasionally the next spiral row of smaller spines is visible on the pen- ultimate whorl covered by the thin upper edge of the former parietal callus. Body whorl large with a series of subequidistant ribs, generally narrow, triangular in cross section, but occa- sionally thickened, especially towards the outer lip; the upper portion of the rib marked by a series of three to four spirally aligned triangular spines, the uppermost one, below the subsutural ramp, the largest, while the third one below the suture is next in size and often marks a distinct angu- losity of the body whorl; ribs marked on the crest with a narrow interrupted chestnut line; ground color between ribs pale yellowish pink or grayish yellowish pink to pinkish gray or brownish pink with a series of bands of varying width of short, axial, zigzag chestnut lines that occasionally be- come irregularly broadened on the adapertural side of the ribs, especially near the outer lip. Large irregularly shaped blotches of chestnut are present between the ribs on the upper portion of the body whorl. The color markings of the body whorl appear more conspicuously banded within 20-674] November 27, 1973 the aperture. Aperture ovate, moderately narrow above where there is a deep subsutural sinus and effuse below; outer lip only slightly thickened, bluntly denticulate at edge, especially in basal half; inner lip almost straight to gently concave. Parietal wall covered by a thin glaze, with a large brown-chestnut splotch at the junction of colu- mellar and parietal lip and two splotches of vary- ing sizes, one on parietal wall near junction of outer lip, the other between the columellar lip and the siphonal fasciole. Measurements (mm.)— length width no. whorls 92.6 63.7 7 large; Carmen Id., Baja Calif. 78.9 48.8 64 medium; Gulf of California 1333 47.9 713 medium; Mulege Bay, Baja Calif. 64.5 39.9 medium; Gulf of California 34.0 21.7 64% small; Gulf of California Synonymy— 1822 Harpa crenata Swainson, Catalogue Shell Colln. Bligh, Appendix, p. 5. (no locality given; Acapulco, Guerrero, Mexico, designated as type locality by Emerson, 1964); 1843, Reeve, Conch. Icon., vol. 1, Harpa, pl. 4, fig. 9a, b, c; 1964, Emerson, American Mus. Novitates, no. 2202, pp. 3-5, fig. 1. 1832 Harpa scriba Valenciennes, in Humboldt and Bonpland, Voyage reg. equinox. Nouv. Continent, pt. 2, Rec. Ob- serv. zool. anat. comp., vol. 2, p. 323 (Acapulco, Mex- ico). 1834 Harpa_ rivoliana Lesson, Illustr. de Zoologie, (12), pl. 36, fig. 1, 2. (Japonia?”); 1860, Sowerby II, The- saurus Conch., vol. 3, p. 171, pl. 232, fig. 12, 13 (Aca- pulco). 1835 Harpa rosea Kiener, Spec. Gen. Icon. Coquilles Viv., vol. 8, Genre Harpe, pp. 11-12 (in part), pl. 5, fig. 8. Not Harpa rosea Lam. 1839 Harpa rosea crenata Gray, Zoology Capt. Beechey’s Voyage, p. 122, pl. 34, fig. 5. Types—The type of crenata Swainson was in Mrs. Bligh’s collection which was sold at auction in May 1822. Although many of the rarities came to the British Museum with the Broderip Collec- tion, no specimen that might be considered to be the type was found in that collection; the type must be considered to be lost. The types of scriba Valenciennes and rivoliana Lesson are not in the Museum National d'Histoire Naturelle in Paris, and their present whereabouts are unknown. Records—MEXICO-BAJA CALIFORNIA: = Mulege Bay (USNM); Loreto; Magdalena Bay (both MCZ); La Paz (AMNH, ANSP, MCZ, RNHL); Cabo San Lucas (USNM). SONORA: Guaymas; Mazatlan (both ANSP, BM, MCZ, USNM). OAX- ACA: Salina Cruz (USNM). GUATEMALA: off Puerto San José (ANSP). COSTA RICA: Salinas Bay (ANSP, BM); Bahia Huevos, W of Puerto Culebra (ANSP); Golfo de Nicoya (USNM). PANAMA: Isla Coiba (Univ. Panama); Isla Pedro Gonzalez, Islas Perlas (ANSP). COLOMBIA: Isla Gorgona (AMNH). Fossil Records—PLEISTOCENE: Punta Coyote, Baja Cali- fornia, Mexico (Hertlein, 1957, p. 59); Rio Colotepec, Oaxaca, Mexico (Palmer and Hertlein, 1936, p. 68). INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Harpa = 259 Harpa myrmia Olsson, 1931 (PI. 229) Range—Lower Oligocene of Peru (Chira forma- tion). Remarks—The few, heavy ribs and more or less angulate shoulder on the body whorl distinguish this Harpa from other species. The suture is cov- ered by the appressed ends of the posterior ends of ribs, and the columellar and parietal callus is thin and spread to some extent over the body whorl. This species is therefore a true Harpa, and the earliest known species of this genus. Through the cooperation of Dr. Katherine V. W. Palmer I have been able to examine the holotype, and have based the following description on this unique specimen. Description—Shell rather small, 32 mm. (1% inches) in length, broadly and angulately ovate, spire broadly conical. Nuclear whorls lost, re- maining whorls 4%. The antepenultimate whorl gently convex, with low widely separated ribs and a few fine spiral striae in the upper part of the interspaces. On the last half of the penultimate whorl (the surface of the earlier part is destroyed) there are two or three fine axial riblets (? growth lines) in the interspaces between the ribs which are slightly angulate at the shoulder; spiral striae are evident in the interspaces, and the lower third to a half of the whorl is covered by the adnate forward-curving upper ends of the ribs of the body whorl. The body whorl has nine strong ribs of which the last three are broader than the others; at the edge of the declivous subsutural ramp the ribs are angulate with an obtuse spine which is particularly apparent on the last four ribs; another pronounced angle is present below, giving the Plate 229. Harpa myrmia Olsson, 1931. Lower Oligocene of Peru. 32 mim. (from Olsson, pl. 20, fig. 7). [20-675] 260 Harpa shell a strongly shouldered appearance; the inter- spaces show again several strong axial striae, crossed by some more or less obscure spiral striae. Outer lip lost; ventral surface covered by a thin callus. Siphonal fasciole strong, lower end broken. Length 31.9 mm., width 23.0 mm. Pal. Res. Inst. No. 2138, Chira formation, near Quercotilla, Chira valley, northern Peru. Synonymy— 1931 Harpa myrmia Olsson, Bull. American Paleontology, vol. 17, no. 63, p. 114, pl. 20, fig. 7. Harpa species A portion of a shell, found in the Lau Islands, eastern Fiji, is noted here in order to call atten- tion to the presence of this genus in Lower Mio- cene times in eastern Melanesia. Although the ventral portion is missing, and the remaining por- tions of the penultimate and antepenultimate whorls are somewhat corroded, it appears that the H. A. Rehder Harpidae expanded portion of the upper ends of the ribs cover in some places about half of that part of the spire whorls between the shoulder angle and the suture. The ribs are fairly closely spaced, and are rather strongly angulate and sub spinose at the shoulder. I am therefore referring this specimen to the Harpa, making this one of the oldest repre- sentatives of the genus. Judging from the size of the fragment, the com- plete shell would measure about 40 mm. in length, close to the maximum size known in Eocithara. It is larger, with a more rounded, less angulate shoulder, than the Indonesian E. muticaeformis Martin of the same age. The shell was collected in tuffaceous limestone on the coast between Tumbou and Tarakua-wai, Lakemba, Lau Islands, Fiji (H. S. Ladd, collector, Sta. L. 389). This is assigned to the base of the Futuna limestone and falls in stage f of the Lower Miocene (Ladd and Hoffmeister, 1945, pp. 25, 99, and personal communication). [20-676] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Austroharpa 261 Genus Austroharpa Finlay, 1931 This genus comprises a series of relatively small recent and fossil species from Australia. The adult shells range in size from 20 to 50 mm. in length, and are characterized by a large paucispiral bulb- ous or dome-shaped protoconch, and axial ribs whose upper ends are only slightly curved for- ward and hardly visible at the suture. The parietal callus is small but distinct and conspicuously mar- gined, though the outer edge may not be raised or thickened. I am dividing this genus into two subgenera, Austroharpa s.s. and Palamharpa Iredale, 1931, largely on the basis of the protoconch, that of the type species of Austroharpa, A. pulligera (Tate, 1889), being larger, bulbous and apparently tilted, while the species of Palamharpa have a smaller, dome-shaped nucleus with the suture marking the earliest whorl being horizontal. In addition A. pul- ligera is larger, 50 mm. in length, while most of the species of Palamharpa do not, to my knowl- edge, reach 40 mm. in length. Subgenus Austroharpa Finlay, 1931 Type: Harpa pulligera Tate, 1889 This Middle Miocene subgenus contains only the single species Austroharpa pulligera (Tate, 1889) and is characterized, as mentioned above, by the large bulbous nucleus which seems to be tilted and quite different in appearance from the smaller, evenly dome-shaped protoconch of the species I am placing under Palamharpa. Because of this striking character and relatively larger shell-size I am inclined to keep it distinct, espe- cially since it was apparently living with a species, of the subgenus Palamharpa, Austroharpa (Pal- amharpa) spirata (Tate, 1889), both being found together in the same Balcombe Clay at Balcombe Bay, Victoria. This suggests that we are dealing with two distinct stocks, whose phylogenetic re- lationship can only be elucidated by the future discovery of related forms. Synonymy— 1931 Austroharpa Finlay, Trans. New Zealand Inst., vol. 62, pt. 1 (May 31), p. 13. Austroharpa pulligera (Tate, 1889) (Pls. 231, 232) Range—Middle Miocene (Balcombian) of Vic- toria. Remarks—This is apparently a rare species, dis- tinct by its size, relatively large among the species of this genus Austroharpa in length, its cassid-like form, and large, bulbous protoconch. The type, from Schnapper Point, near Mornington, Victoria, north of Balcombe Bay, measures 50 mm_ in length; Dr. Thomas A. Darragh (in litt.) informs me that the National Museum of Victoria has eight specimens from Balcombe Bay ranging in length from 38 to 46 mm. He states that the spe- cies occurs also at Muddy Creek, near Hamilton, Victoria. As I have not seen specimens I am copy- ing Tate’s description: Shell thin, oval, with a rather short spire, ending in a very large hemispheric pullus, with the tip laterally immersed; the second turn of the pullus almost concealed by the first ordinary whorl. Or- dinary whorls one and a half, subangulated; orna- mented with thin, slightly elevated lamellae, Plate 230. Enlarged protoconchs of species of Austroharpa (Palamharpa). Fig. 1. A. (P.) punctata (Verco). Off Venus Bay, South Australia, USNM 706971. Fig. 2. A. (P.) exquisita (Ire- dale). off Burleigh Heads, Queensland, ANSP 314410. Fig. 3. A. (P.) sulcosa (Tate). Miocene, Hamilton, Victoria, USNM 157219. (all X 10). [20-685] 262. Austroharpa H. A. Rehder Harpidae £ F ee we he Plate 231. Austroharpa pulligera (Tate). Holotype, 50 mm. in length. South Australian Museum, Tate Colln. 703. which are vaulted on the angulation. Last whorl oval-oblong, somewhat ventricose over the suture, ornamented with about 25 thin, slightly elevated lamellae, which are raised into vaulted scales on the shoulder; the interspaces with coarse axial [20-686] ade Yu \ \, Plate 232. Protoconch of Austroharpa (Austroharpa) pulligera (Tate, 1589). (from Cotton and Woods, 1933, p. 46, fig. 8) X 7.5. striae; base spirally wrinkled. Aperture narrow- oval; outer lip slightly ascending on the penulti- mate whorl, its margin much thickened. Dimensions (in mm.)—length, 50, breadth 30, length of aperture 42, diameter of pullus 4.5. Synonymy— 1889 Harpa pulligera Tate, Trans. Proc. Rep. Royal Soc. South Australia, vol. 11, p. 151, pl. 6, fig. 9 (Blue clays at Schnapper Point, Mornington, Victoria). 1913 Harpa (Eocithara) pulligera Tate, Verco, Trans. Royal Soc. South Australia, vol. 37, p. 447. 1931 Austroharpa pulligera Tate, Finlay, Trans. New Zea- land Inst., vol. 62, p. 13. 1931 Deniharpa pulligera Tate, Iredale, Rec. Australian Mus., vol. 18, p. 230. November 27, 1973 Subgenus Palamharpa Iredale, 1931 Type: Palamharpa exquisita Iredale, 1931 This group of recent and fossil Australian spe- cies is characterized by its rounded, dome-shaped paucispiral protoconch (1% whorls), generally small size—from 20 to 35 mm. (the exception is a specimen reconstructed from a fragment of Au- stroharpa (Palamharpa) loisae Rehder which measures 45.8 mm. in length). In sculpture the species are very variable ranging from those with rather crowded lamellar ribs and cancellate sculp- ture through ones with very little or no spiral sculpture to a smooth species with only weakly indicated, widely separated, axial ribs. The recent species have a rather distinct notch at the upper end of the outer lip below its junction with the su- ture, which is not apparent in the fossil species. This subgenus has been considered by some (Cotton and Woods, 1933, p. 47; Wenz, 1943, p. 1310) to be a synonym of Austroharpa s.s., but I believe that the striking difference in the proto- conch is sufficiently important to warrant for the present their separation. In the many specimens of Harpidae, recent and fossil, that I have exa- mined I have found the protoconchs to be quite constant. The living species of the subgenus Palamharpa are found in moderately deep water from southern Queensland southward around the southern Au- stralian coast to off Perth, Western Australia. The fossil species range from Upper Oligocene to Upper Pliocene. Synonymy— 1931 Palamharpa Iredale, Rec. Australian Museum, vol. 18, no. 4 (June 29), pp. 230, 233 (type, by original desig- nation: Palamharpa exquisita Iredale). 1931 Deniharpa Iredale, ibid. (Type, by original designation: Harpa clathrata Tate). 1931 Trameharpa Iredale, ibid. (type, by original designation: Harpa spirata Yate). Austroharpa exquisita (Iredale, 1931) (PI. 230, fig. 2; Pls. 233, 237, figs. 4, 5) Range—From off Burleigh Heads, southern Queensland, to Bass Straits, Victoria and Tas- mania, in 25 to 80 fathoms. INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Austroharpa 263 Remarks—This small, deepwater species was once considered, after A. (P.) punctata (Verco), the rarest of the harp shells. It is still uncommon in collections but has been brought up in recent years from moderately deep waters off southern Plate 233. Austroharpa (Palamharpa) exquisita (Iredale). Holotype, Australian Museum no. C. 57753, from off Twofold Bay, New South Wales, Australia. [20-689] 264. Palamharpa New South Wales and eastern Victoria by com- mercial fishermen; in the latter locality, according to Mrs. M. C. Griffiths of Lakes Entrance, Victoria, about 10 specimens have been found in the last thirteen years. Habitat—According to Mrs. M. C. Griffiths (in litt.), specimens of this species are dredged in Bass Straits in an area about 7-9 miles ESE of Lakes Entrance, Victoria, on a bottom consisting of dead shell debris that formed almost a coarse shell sand. Description—Shell small, 24-29 mm. in length, broadly ovate with a conical, turrited spire. Proto- conch dome-shaped, of 1% smooth whorls, the earliest part of the first whorl low and well im- mersed below the horizontal suture in succeeding whorl, last part more convex and impressed at suture; postnuclear whorls of spire with an in- creasingly angled shoulder, weak distant ribs, and broad low, obscure ridges, one in subsutural ramp, one at shoulder and two below the shoulder; the angulate shoulder and flattened subsutural ramp gives the spire a turrited appearance; be- tween the ribs which become increasingly sub- lamellar, are numerous axial ridges, about 3 to 6. Body whorl strongly angulate, with about 22 to 27 low, sublamellar axial riblets which are rendered weakly scalloped by the broad, rounded spiral ridges they cross; these spiral ridges number 10-12 below the shoulder; the fine axial ridges between the ribs are sharply and finely sublamellar. Color of nucleus yellowish-pink, of body whorl from pale yellowish pink or moderate yellowish pink to dark orange yellow, with splotches of moderate reddish orange or grayish reddish orange on the subsutural ramp or arranged in obscure bands on body whorl, the spots darker on the lamellar ribs where they cross the spiral ridges. Aperture elongate, inner edge rather straight; outer lip gently arcuate, slightly flattened in center, somewhat flaring, thickened externally, with a definite notch below its junction with the body whorl; parietal-colu- mellar junction indistinctly angulate, parietal cal- lus narrow, distinctly marginate, raised in the area of the base of the body whorl and the siphonal fasciole; siphonal canal rather deep and directed upward. The soft parts of a specimen from Twofold Bay, New South Wales, sent to me by Dr. D. F. McMi- chael, then at the Australian Museum, Sydney, preserved in alcohol for some time, show an ani- mal of which the propodium and head and anterior part of the metapodium are without spots; the posterior portion shows spots which are densest at the posterior end. H. A. Rehder Harpidae Measurements (mm.)— no. whorls 24 15 5 length width Holotype, Twofold Bay, N.S.W. 28.4 17 5+ Burleigh Heads, Queensland 29.0 17.3 5 Lakes Entrance, Victoria O17 12.5 5 Eden, New South Wales Synonymy— 1931 Palamharpa exquisita Iredale, Rec. Australian Mus., vol. 18, no. 4 (June 29), p. 230, pl. 22, fig. 8. 1933 Austroharpa exquisita Iredale, Cotton and Woods, Rec. South Australian Mus., vol. 5, p. 47; 1962 Macpher- son and Gabriel, Marine Molluscs of Victoria, p. 215, figure 257; 1971 Wilson and Gillett, Australian Shells, p. 110, pl. 72, fig. 6. Types—The holotype is in the Australian Mu- seum, No. C. 57753, and the type locality, which was not mentioned by Iredale, is off Twofold Bay, New South Wales, in 45 fms. Records—QUEENSLAND: off Burleigh Heads, in 34 fms. (ANSP). NEW SOUTH WALES: 11 mi W of Crowdy Head, in 50 fms. (AMS): E of Sidney, in 40-82 fms. (AMS); off Crook- haven Bight, in 30-35 fms. (Colln. W. A. Trenerry); Ulladulla (Colln. G. Thornley); off Crabo Id., Twofold Bay, in 50 fms. (AMS); off Twofold Bay, in 45 fms. (AMS, NMV); off Eden, in 50-60 fms. (USNM). VICTORIA: off Hospital Creek, in 30-60 fms.; ESE of Lakes Entrance, in 21-26 fms. (both Colln. M. C. Griffiths); off Lake Tyers (Collns. C. J. Gabriel, W. S. Ayres). TASMANIA: off Deal Id., Kent Group, Bass Straits, in 33 fms. (Garrard, 1961). Austroharpa loisae Rehder, new species (Pl. 237, figs. 3, 6) Range—From WSW of Cape Naturaliste to NW of Rottnest Island, Western Australia. Remarks—tThis strikingly sculptured species is most closely related to A. (P.) exquisita (Iredale) from the southeastern coast of Australia. It differs, however, in being more slender, not as strongly shouldered, in the axial and spiral sculpture being stronger and more regular, and in the protoconch and early whorls being lemon yellow rather than pinkish yellow in color. A fragment of a large specimen found in 80 fathoms NW of Rottnest Island (SAM 34-70) is the basis for the maximum size mentioned in the description and listed in the measurements below. The height of the penultimate whorl, measured from suture to suture, was compared with the same measurement taken from the holotype; the relationship between these two measurements was equated with the total length of the holotype, and by this means an estimated length for the large specimen was determined. The holotype has been figured by Wilson and Gillett in their book “Australian Shells” (1971, pl. 72, fig. 6a) as Austroharpa exquisita Iredale, | 20-690 | November 27, 1973 Plate 234. Geographic distribution of: 1, Austroharpa (Palam- harpa) exquisita (Iredale), and 2, Austroharpa (Palamharpa) loisae Rehder, new species. who noted that this form may prove to be a dis- tinct subspecies or species. This beautiful species is named for my wife in appreciation of her ever willing assistance, en- couragement, and understanding. Habitat—This species has been dredged from depths of from 70 to 103 fathoms, on sandy bot- tom with sponge and bryozoa. Description—Shell of medium to relatively large size, thin, adults measuring from 28.4 to 45.8 mm. (1% to 1% inches), ovate, with a rather elevated, conical, and turrited spire. Protoconch almost hemispherical, dome-shaped, paucispiral, of 12+ smooth whorls, moderate yellow in color. Post- nuclear whorls 3% in number, the obtuse shoulder marked by a spiral cord, in addition to which there is an obscure spiral cord on the subsutural ramp and more pronounced equidistant spiral cords below the shoulder (3 on the antepenultimate whorl, and 4 on the penultimate whorl). Crossing these cords are thin, rather distant, raised lamel- lar ribs (25 in the penultimate and 29 on the last whorl); these ribs are noticeably scallopped where they cross the equidistant spiral cords, the scallops being highest on the shoulder and subsutural cords; if the body whorl is viewed against a light, the axial ribs and cords make a very regular, re- ticulated pattern; between the axial ribs are 6 to 10 very fine, rather regular and somewhat sepa- rated, lamellar axial riblets. Color is a pinkish white or yellowish white to a yellowish gray, with the protoconch and first postnuclear whorls a moderate yellow; there are numerous small red- dish brown spots where the ribs cross the cords, most noticeable on the abapertural side of the lamellar ribs, and occasionally a few larger spots of pale reddish brown on the subsutural ramp. INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Austroharpa 265 Aperture elongate, semilunate, with inner lip only slightly angled at juncture of parietal and colu- mellar portions, and outer lip gently arcuate, somewhat broadly reflected and a little thickened internally, a small but distinct notch present at upper end below junction with body whorl; parie- tal callus thin, allowing sculpture underneath to be apparent, well margined, with a pronounced margin in adult shells; anteriorly, in the region of the well-developed and_ strongly lamellate _ si- phonal fasciole, the margin is suberect, resulting in a noticeable pseudumbilical chink; anterior si- phon rather deep, upturned. Measurements (mm.)— length width no. whorls 28.9 16.7 5 Holotype 28.4 16.5 4% ~~ Paratype No. 1 45.8° _— _— Paratype No. 3 36.4° - — Paratype No. 4 “length (approximate) computed from height of penultimate whorl. Synonymy— 1971 Austroharpa exquisita Iredale, Wilson and Gillett, Au- stralian Shells, p. 110 (in part), pl. 72, fig. 6a. Not Palamharpa exquisita Iredale, 1931. Types and Records—WESTERN AUSTRALIA: WSW of Cape Naturaliste, in 75 fms., broken shell (paratype no. 4, WAM 129-63); W of Rottnest Id., in 60 fms. (WAM 156-72); NW of Rottnest Id., in 70 fms. (holotype, WAM 1784-69); NW of Rottnest Id., in 100-103 fms. (paratype no. 1, WAM 31-64); NW of Rottnest Id., in 85 to 95 fms. (paratype no. 2 and frag- ment, WAM) 127/128-63); NW of Rottnest Id., in SO fms. (fragment, paratype no. 3, WAM 34-70); WSW > of Dongara, in 60 fms. (paratype, USNM 707703; WAM 158-72; WAM 159-72); NW of Bluff Point, in 60 fms. (WAM 157-72). Austroharpa punctata (Verco, 1896) (PI. 230, fig. 1; Pls. 235, 236) Range—The eastern half of the Great Australian Bight, South Australia, from Nuyts Archipelago to Encounter Bay. Remarks—tThis striking shell was the first of the living members of this genus to be described, and is still one of the rarest and most sought-after shells; only about twenty specimens are known. It is characterized by its size, relatively large for the subgenus, its inflated shape, and its smooth col- ored shell which is without any obvious spiral sculpture and has only low, obscure varices with a groove immediately in front of them and marked by scales on the subsutural ramp. It is quite dis- tinct from any other known species, its closest relative being the smaller, more strongly sculp- tured species A. (P.) wilsoni Rehder described below. [20-691 | 266 = Palamharpa H. A. Rehder Harpidae Plate 235. Austroharpa (Palamharpa) punctata (Verco). Holo- type. South Australian Museum D. 13516. Off Newland Head, Encounter Bay, South Australia, in 20 fathoms Description—Shell moderately large, 32 to 36.3 mm. in length, inflated-ovate, with short, broadly conical spire, the whorls of which are flattened subsuturally and are convex below the rounded, not angulate shoulder. Protoconch rather large, hemispherical, dome-shaped, of 1% smooth whorls, the initial 14 whorls microscopically granulose. Postnuclear whorls 2% in number, convex, with a flat subsutural shelf, marked by low axial varices which gradually increase in strength; the varices are the outer edges of former lips, with the suc- ceeding shell growth starting below the level of that non-reflected lip-edge, forming in this way a series of distant, overhanging steps, which are highest near the shoulder and gradually diminish in height towards the base; above the rounded shoulder and on the subsutural shelf the varices are marked by large, erect, forward-leaning, con- cave scales; between the varices the shell is smooth except for irregular, microscopic wavy striae. Color of protoconch and spire whorls a moderate yellowish orange-pink, body whorl vary- ing from a moderate or strong yellowish pink to occasionally a deep yellowish pink or salmon color, with obscure bands and irregular spots of white; the darker, strong yellowish pink color may be patterned as large spots arranged in three bands—on the subsutural shelf and on the middle and lower half of the body whorl; the whole shell [20-692] is usually marked with irregular flecks and spots of various shades of reddish brown, the spots of various sizes and shapes but most frequently tri- angular, elongate, or sagittate; specimens are OC- casionally found without spots. Aperture elongate, semilunate, outer lip gently arcuate, rather broadly reflected, only slightly thickened externally and internally, inner surface showing the pink colora- tion of the external, obscure, darker banding; a strong triangular notch is present at the upper end, at the junction with the parietal wall. Parietal cal- lus small, very thin, with the outer margin ob- scure; columellar callus with a conspicuous, thick- ened margin at the siphonal fasciole; siphonal canal short, deep, upturned. According to a note by the collector of a living specimen from Thorny Passage, the animal is pale orange. The dried soft parts extracted from a specimen from Venus Bay (USNM 706971), and softened in Aerosol O.T., show scattered spots of reddish brown on a pale ground on both foot and tentacles; the siphon is also spotted and _ indis- tinctly annulate. Measurements (mm.)— length width no. whorls 34.0 22.0) 4% Holotype, SAM D13516 32.5 21.5 de Paratype, SAM D459 35.0 24.0 4s SAM D460 34.5 24.9 4s USNM 706971 34.7 25.1 4% = Colln. Hurrell 36.4 24.9 4 Colln. Delaney 31.9 21.4 43 Colln. Delaney November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Austroharpa 267 Plate 236. Austroharpa (Palamharpa) punctata (Verco). Left fig. off Venus Bay, South Australia, USNM 706971. Center fig. off Encounter Bay, South Australia, Colln. S. T. Delaney. Right fig. off Venus Bay, South Australia, Colln. D. Hurrell. (all natural size). Synonymy— 1896 Harpa punctata Verco, Trans. Royal Soc. South Austra- lia, vol. 20, p. 218, pl. 6, figs. 3, 3a, 3b (Newland Head, South Australia). 1913 Harpa (Eocithara) punctata Verco, Trans. Royal Soc. South Australia, vol. 37, pp. 446-447. 1931 Austroharpa punctata Verco, Finlay, Trans. New Zea- land Inst., vol. 62, p. 13; 1933, Cotton and Woods, Rec. South Australian Museum, vol. 5, p. 47; 1971, Wilson and Gillett, Australian Shells, p. 110, pl. 72, figs. 5, 5a Types—The holotype (SAM D13516) and para- type (SAM D459) are in the South Australian Mu- seum. The type locality is off Newland Head, at the northern end of Encounter Bay, South Austra- lia, in 20 fathoms. Records—SOUTH AUSTRALIA: St. Francis Id., Nuyts Ar- chipelago (SAM D460); off Venus Bay, in 27 ims. (USNM, Colln. D. Hurrell); near Port Lincoln (fide H. M. Laws); Thorny Passage, in 65 to 70 ft. (CollIn. S. T. Delaney); Emu Bay, Kan- garoo Id. (Colln. F. L. Saunders, fide H. M. Laws); Americarr River, Kangaroo Id. (teste Verco, 1913); Backstairs Passage, in 22 fms. (teste Verco, 1896); Normanville (SAM); off New- land Head, Encounter Bay, in 20 fms. (holotype, SAM); En- counter Bay, in 20 fms. (SAM, Colln. S. T. Delaney). Austroharpa wilsoni Rehder, new species (PI. 237, figs. 1, 2) Range—From off Cape Leeuwin to off Dongara, Western Australia. Remarks—This species is most closely related to A. (P.) punctata Verco by reason of its similar sculpture; it is, however smaller, more slender, with a relatively higher spire, and the shell a uni- form whitish yellow with only occasional spots on the subsutural shoulder. It is named for Barry R. Wilson of the Western Australian Museum, through whose generosity I was able to study the Harpidae of Western Au- stralia. Habitat—All specimens were dredged in depths of from 60 to 120 fathoms on a sandy bottom, of- ten with sponges, bryozoa, and starfish. Description—Shell small, from about 20 to 25.7 mm. in length, thin, ovate, with an elevated coni- cal spire. Protoconch, hemispherical, dome- shaped, of 1% whorls that appear smooth but are microscopically — granulose; convex, with a very weakly angulate shoulder, and a slanted subsutural ramp; the sculpture consists of distant axial ribs, and in the earliest whorls a few low and broad spiral cords that gradually be- come obscure and are only very weakly indicated on the body whorl at and below the shoulder; in the early whorl the axial varices are more erect and rib-like, but later the varices are lower and have the appearance of low slightly overhanging postnuclear whorls steps; on the subsutural ramp the varices are more elevated and vaulted and occasionally form an erect, hollow triangular scale. The whole surface is superficially smooth but shows under highpower magnification very fine, obscure, wavy striae and coarser and irregular axial growth wrinkles. Color grayish yellow, occasionally with pale orange brown splotches on the shoulder and series of small light reddish brown spots on the varices be- hind the sharp edge; these seem to be arranged in spiral series, and on fresh shells a faint indication of spiral banding can be seen on the body whorl; the fine edge of the body-whorl varices shows a reddish brown color where these bands cross, and on the thickened edge of the outer lip the series of red brown blotches is conspicuous; these spots are continued on the inner edge of the thickened outer lip where in some instances the spots become a dark pink color; in dead shells the spots on the lip may disappear; in the holotype the siphonal fas- ciole is flushed with pink and the columellar callus [20-693] 268 Palamharpa H. A. Rehder Harpidae where it crosses the fasciole is a pale pink color. Aperture elongate-semilunate with outer lip gently arcuate, thickened internally, narrowly reflected, with a conspicuous, moderately deep sinus below its juncture with the parietal wall; parietal callus small, thin, the margin low but definite on the parietal wall, thickened on the columellar portion with narrow chinks above and below where it crosses the siphonal fasciole; siphon deep, up- turned, its inner surface with a pale rosy flush. Wil- son (in litt.) describes the living animal as “white with sparse lemon yellow spots on the sides of the foot and on the head, eye stalks lemon yellow, penis large and white.” Measurements (mm.)— length width no. whorls 25.3 14.7 5+ Holotype = 36-70) 25.7 15.6 4 Paratype No. 1 (WAM 125-63) 24.4 15.0 ao Paratype No. 2 (USNM 703249) 22.8 14.1 de Paratype No. 3 (USNM 703250) 21.6 12.5 4 Paratype No. 4 (WAM) 19.7 11.5 4% Paratype No. 5 (WAM 35-70) Types—The type locality is NW of Rottnest Is- land, off Perth, Western Australia in 80 fathoms on a bottom of sand with bryozoa and sponges; collected on a cruise of the “Bluefin,” Sept. 15, 1965. The holotype is WAM 36-70. Records—WESTERN AUSTRALIA: W of Cape Leeuwin, in 76-80 fms. (WAM 153-72); WNW of Cape Freycinet, in 107- 129 fms. (WAM 154-72); W of Cape Naturaliste, in 96-100 fms. (WAM 155-72); SW of Garden Id., in 81-84 fms. (WAM 152-72); W of Rottnest Id., in 75 fms. (WAM 38-70); WNW of Rottnest Id., in 95-96 fms. (USNM 703250; WAM _ 122/124- 63); NW of Rottnest Id., in 70-103 fms. (USNM 703249; WAM 35-70; 36-70; 125/126-63); WSW of Dongara, in 60 fms. (WAM 150-72); W of Dongara, in 80 fms. (WAM 151-72). *\ | 2\ & (3 Plate 238. Geographic distribution of: 1, Austroharpa (Palam- harpa) punctata (Verco), and 2, Austroharpa (Palamharpa) wilsoni Rehder, new species. Plate 237. Figs. 1, 2. Austroharpa (Palamharpa) wilsoni Reh- der, new species. 1, holotype, WAM 36-70. 2, paratype, WAM 123-62. Figs. 3, 6. Austroharpa (Palamharpa) loisae Rehder, new species. 3, holotype, WAM 1754-69. 6, paratype, WAM 31-64. Figs. 4, 5. Austroharpa (Palamharpa) exquisita Iredale. 4, off Burleigh Heads, Queensland, ANSP 314410. 5, off Eden, New South Wales, USNM 634267. (all natural size). [20-694] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Austroharpa 269 Austroharpa sulcosa (Tate, 1889) (Pl. 239) Range—Middle Miocene (Balcombian) of Vic- toria. Remarks—A species that seems closest to A. (P.) exquisita Iredale but with a lower spire, a some- what more broadly ovate shape, and with stronger axial and spiral sculpture. The whorls are actually subsuturally canaliculate by the raised angular shoulder on which the axial ribs form erect lamel- lar and triangular scales. The parietal callus is thin judging from the single specimen with underde- veloped outer lip from the type locality I have been able to examine personally (USNM_ 157219); this specimen measures 28.2 mm. in length. According to N. H. Ludbrook, this species is also found in the Fyansford Clay of the Balcombian at Shelford, Victoria. Synonymy— 1889 Harpa sulcosa Tate, Trans. Proc. Rep. Royal Soc. South Australia. vol. 11, p. 150, pl. 6, fig. 10 (Muddy Creek, Hamilton, Victoria). 1897 Harpa (Eocithara) sulcosa Tate, Harris, Cat. Tert. Moll. Dept. Geol. British Museum, pt. 1, p. 79. 1931 Austroharpa sulcosa Tate, Finlay, Trans. New Zealand Inst., vol. 62, p. 13; 1933 Cotton and Woods, Rec. South Australian Mus., vol. 5, p. 47, fig. 2 (protoconch). 1931 Refluharpa sulcosa Yate, Iredale, Rec. Australian Mu- seum, vol. 18, p. 230. Nome Harpa sulcosa, spec. nov. Plat Hlah. Coecene. Muddy Cy eet Plate 239. Austroharpa (Palamharpa) sulcosa (Vate). Holo- type (28.2 mm. in length) and paratypes. South Australian Museum, Tate Colln. 718. Austroharpa tatei Finlay, 1931 (Pl. 240) Range—Pliocene (Dry Creek Sands) near Ade- laide, South Australia. Remarks—This species is close to A. (P.) sulcosa as Finlay and Ludbrook point out, and even more closely related to A. (P.) loisae Rehder of which it may represent an ancestral form. In number of lamellate axial ribs on the body whorl (33) it is intermediate between sulcosa, which has about 38 and loisae, which has 29. It is less strongly angulate at the shoulder than sulcosa, lacks the spines at the shoulder, and the spiral sculpture is stronger. In all these features it is close to A. (P.) loisae. The type measures 25.5 mm. in length and 17 mm. in diameter. It is the Finlay Collection (no. 67) in the Auckland Museum, New Zealand. Synonymy— 1931 Austroharpa tatei Finlay, Trans. New Zealand Inst., vol. 62, p. 14 (“Older Pliocene,” Abbatoirs Bore, Adelaide, South Australia). 1958 Harpa (Austroharpa) tatei Finlay, Ludbrook, Trans, Royal Soc. South Australia, vol. 81, p. 73, pl. 4, fig. 5. Plate 240. Austroharpa (Palamharpa) tatei Finlay. Holotype, Auckland Institute and Museum. 25.5 mm. (copied from Lud- brook, 1958). Austroharpa spirata (Tate, 1889) (PI. 241) Range—Middle Miocene (Balcombian) of Vic- toria. Remarks—This species is closely related to sul- cosa Tate but has stronger sculpture, with the axial ribs broader and subequal and with more pronounced spiral cords, both resulting in a strongly fenestrated sculpture. Below the narrow Plate 241. Austroharpa (Palamharpa) spirata (Yate). Shelford, Victoria. 26.5 mm. in length. South Australian Museum, P4257 [20-695] 270 = Palamharpa subsutural channel the subsutural ramp slants to the angulate shoulder, with the axial ribs bearing erect scales at the border of the narrow sutural channel. The holotype of A. (P.) spirata, which measured 35 mm. in length has disappeared, according to N. H. Ludbrook, who has furnished the photo- graph which depicts a specimen from Shelford, Victoria. Synonymy— 1889 Harpa spirata Tate, Trans. Proc. Rep. Royal Soc. South Australia, vol. 11, p. 150, pl. 6, fig. 3 (Blue clays at Schnapper Point—Mornington, Victoria). 1931 Austroharpa spirata Tate, Finlay, Trans. New Zealand Inst., vol. 62, p. 13; 1933, Cotton and Woods, Rec. South Australian Mus., vol. 5, pp. 45, 47, fig. 3 (proto- conch). 1931 Trameharpa spirata Tate, Iredale, Rec Australian Mu- seum, vol. 18, p. 230. Austroharpa tenuis (Tate, 1889) (Pls. 242, 243) Range—Lower to Middle Miocene (Batesfordian to Balcombian) of Victoria. Remarks—This species is larger than most of the other species and differs from the previous two species in the reduction of the spiral sculpture to low, more or less obscure ridges. It resembles in this respect the recent A. (P.) exquisita Iredale but Le aD ~ LD Plate 242. Austroharpa (Palamharpa) tenuis (Tate). Holotype (34.5 mm. in length) and paratypes. South Australian Museum, Tate Colln. 702 H. A. Rehder Harpidae the latter has a higher spire, more angulately shouldered whorls, and the ribs are not spinose at the shoulder. Besides the type locality of Muddy Creek, Ham- ilton, Victoria, this species is also found at Royal Park, Victoria in the Newport formation of the Balcombian (Middle Miocene), and from Flinders, Victoria, in the Batesfordian of the Lower Mio- cene. The type according to Tate, measured 36 mm. in length and 23 mm. in width. A specimen in the collections of the USNM measures 36.2 in length and 21.7 mm. in width. Synonymy— 1889 Harpa tenuis Tate, Trans. Proc. Rep. Royal Soc. South Australia, vol. 11, p. 151, pl. 6, fig. 1 (Muddy Creek, Hamilton, Victoria). 1897 Harpa (Eocithara) tenuis Tate, Harris, Cat. Tert. Moll. Dept. Geol. Brit. Mus., pt. 1, p. 80, pl. 4, figs. 4a, 4b (protoconch). 1931 Austroharpa tenuis Tate, Finlay, Trans. New Zealand Inst., vol. 62, p. 13; 1933, Cotton and Woods, Rec. South Australian Museum, vol. 5, p. 47, fig. 9 (proto- conch). 1931 Deniharpa tenuis Tate, Iredale, Rec. Australian) Mu- seum, vol. 18, p. 230. Plate 243. Austroharpa (Palamharpa) tenuis (Tate, 1889). Clifton, Victoria. Balcombian (Middle Miocene). USNM 647308. 36.2 mm. Austroharpa abbreviata (Tate, 1889) (Pl. 244) Range—Middle Miocene (Balcombian) of Vic- toria. Remarks—This species has a close resemblance in shape and nature of axial ribs to A (A.) pulligera Tate, but differs in size and in the nature of the protoconch. It is fairly closely related to A. (P.) tenuis Tate but has fewer axial ribs, the whorls are not as strongly shouldered, the ribs do not bear the erect scales at the shoulder angulation, and the spiral sculpture appears to be absent or is at least very obscure. {20-696 | November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Austroharpa 271 cE TIS Harpa abbreviat tece WAR 7 mudd yech Plate 244. Austroharpa (Palamharpa) abbreviata (Tate). Holotype (27.5 mm. in length). South Australian Museum, Tate Colln. 710. Synonymy— 1889 Harpa abbreviata Tate, Trans. Proc. Rep. Royal Soc. South Australia, vol. 11, p. 150, pl. 6, fig. 7. 1897 Harpa (Eocithara) abbreviata Tate, Harris, Cat. Tert. Moll. Dept. Geol. British Museum, p. 81, pl. 4, figs. 5a- b (protoconch). 1931 Austroharpa abbreviata Finlay, Trans. New Zealand Inst., vol. 62, p. 13; 1933, Cotton and Woods, Rec. South Australian Museum, vol. 5, pp. 45, 47, fig. 7 (protoconch). 1931 Deniharpa abbreviata Tate, Iredale, Rec. Australian Museum, vol. 18, p. 230. Plate 245. Austroharpa (Palamharpa) clathrata (Yate). Holo- type (38.2. mm. in length) and paratypes. South Australian Museum, Tate Colln. 699. Austroharpa clathrata (Tate, 1889) (PI. 245) Range—Lower Miocene (Batesfordian) of Vic- toria. Remarks—tThis relatively large species is distin- guished by the rather broad shell with a low coni- cal spire, the fairly distant, narrow axial ribs crossed by strong subequidistant cords, the inter- sections at and just below the shoulder cord marked by subspinose nodes. The parietal callus is distinctly margined at its outer edge. The holo- type measures about 39 mm. Synonymy— 1889 Harpa clathrata Tate, Trans. Proc. Rep. Royal Soc. South Australia, vol. 11, p. 151, pl. 6, fig. 8 (Murray River cliffs, near Morgan, South Australia). 1931 Austroharpa clathrata Tate, Finlay, Trans. New Zealand Inst., vol. 13, p. 13; 1933, Cotton and Woods, Rec. South Australian Museum, vol. 5, p. 47, fig. 5 (pro- toconch). 1931 Deniharpa clathrata Yate, Iredale, Rec. Australian Mu- seum, vol. 18, p. 230. Austroharpa pachycheila (Tate, 1894) (Pl. 246) Range—Upper Oligocene (Janjukian/Longford- ian) of Victoria. Remarks—This and the following species, A. (P.) cassinoides Tate, were placed by Finlay (1931, p. le, + eiethe. kh EB $ é Es e y%, Name Nakfa pack checle ST ero lluh. Secerre, Spreng € eek (T f] 12 Plate 246. Austroharpa (Palamharpa) pachycheila (Tate). Holotype (at center: 27.5 mm. in length) and paratypes. South Australian Museum, Tate Colln. 712. [20-697] 272 = Palamharpa H. A. Rehder Harpidae 12) in the family Cassidae, “perhaps as Oniscidia,” because of the character of the protoconch, and probably also because of their heavy “cassid” ap- pearance. Three specimens of pachycheila are in the collections of the U.S. National Museum of Natural History, and all have a protoconch that is close to that found in the recent Austroharpa (Pal- amharpa) exquisita Iredale and related species, differing merely in being smaller, with a smaller initial whorl, and a somewhat more impressed su- ture; these appear to be differences of degree only. Drawings of the apical whorls of the holotype and paratypes of pachycheila, kindly sent me by Dr. N. H. Ludbrook, show that both kinds of proto- conch are present in the type lot. A. (P.) pachycheila Tate is a rather stout shell, strongly angulate at the shoulder, the ribs not lamellate but triangular in cross-section; in the first postnuclear whorl spiral cords crossing the axial ribs create a cancellate sculpture; the spiral cords diminish in strength, except for the one at the shoulder angle, particularly in the area below the shoulder. The outer lip is thickened and re- flected, and the parietal callus is usually conspicu- ously marginate. In general aspect and characters this species is close also to A. (P.) clathrata Tate. The holotype measures 27.5 mm. in length. The three specimens mentioned above measure from 23.8 to 26.8 in length, and from 15.0 to 17.4 mm. in width. Synonymy— 1894 Harpa pachycheila Tate, Jour. Royal Society New South Wales, vol. 27, p. 173, pl. 11, fig. 5 (Spring Creek—Tor- quay, Victoria). Plate 247. Austroharpa (Palamharpa) cassinoides (Tate). Holotype, 29 mm. in length. South Australian Museum, Tate Colln. 692. 1931 [Oniscidia] pachycheila Tate, Finlay, Trans. New Zea- land Inst., vol. 62, p. 12. 1931 Deniharpa pachycheila Tate, Iredale, Rec. Australian Museum, vol. 18, p. 230. 1933 Austroharpa pachycheila Tate, Cotton and Woods, Rec. South Australian Museum, vol. 5, pp. 45, 47, fig. 6 (protoconch). Austroharpa cassinoides (Tate, 1889) (Pl. 247) Range—Lower Pliocene (? or Upper Miocene) to Upper Pliocene of New South Wales and South Australia. Remarks—This species, together with A. (P.) pachycheila Tate and possibly also A. (P.) clathrata form a group of rather broad, angulate or subangu- late, stout species with non-lamellar ribs, thick- ened, reflected lip that resemble in general ap- pearance certain members of the Cassidae, and are rather dissimilar to most species of the living Har- pidae. However, because their protoconchs ap- proach those of more typical members of the sub- genus Austroharpa (Palamharpa), and since most Australian workers, who have seen more material than I have, retain these species in this group, I follow their example. This present species is the most aberrant of all members of this subgenus, and is characterized by its short, stout and broad shell, short spire, dimin- ished number of strong non-lamellar ribs that are subnodose at the shoulder, lack of spiral sculpture, and a thickened, reflected outer lip, whose upper end is flexed upwards, projecting over the penul- timate whorl to the antepenultimate whorl. The type measures about 29.5 mm. in length and 22 mm. in width. Synonymy— 1889 Harpa cassinoides Tate. Trans. Proc. Rep. Royal Soc. South Australia, vol. 11, p. 150, pl. 6, fig. 4 (Well sink- ing. Murray Desert—Tareena, New South Wales). 1931 [?Oniscidia] cassinoides Tate, Finlay, Trans. New Zea- land Inst., vol. 62, p. 12. 1931 Deniharpa cassinoides Tate, Iredale, Rec. Australian Museum, vol. 18, p. 230. 1933 Austroharpa cassinoides Tate, Cotton and Woods, Rec. South Australian Museum, vol. 5, p. 47, fig. 4 (proto- conch). 1958 Harpa (Austroharpa) cassinoides Tate, Ludbrook, Trans. Royal Soc. South Australia, vol. 81, p. 74, pl. 4, fig. 4. [20-695] November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 16 Index bo ~l 3 INDEX TO HARPIDAE NAMES IN VOL. 3, NO. 16 Looseleaf subscribers should keep this index at the be- ginning of the family Harpidae. The family begins on page 207 [looseleat p. 20-601]. In this index, the number following the name refers to the pagination found at the top of the page in vol. 3, no. 16. The column at the right is the looseleaf pagination. All new names proposed in this number are in boldface type. [looseleaf } abbreviata Tate, 270 20-697 altavillensis Defrance, 225 20-625 americana Pilsbry, 257 20-673 amoretta Link, 242 20-658 amouretta Réding, 240 20-656 articularis Lamarck, 249 20-665 Austroharpa Finlay, 261 20-6585 bellardii Sacco, 229 20-629 bellardii madachi Noszky, 216 20-610 bibliography, 220 20-618 biology, of Harpa, 209 20-603 birmanica Vredenburg, 228 20-628 brochoni ‘Benoist’ Cossmann, 256 20-672 cabritii Fischer, 252 20-668 californiensis Vokes, 225 20-625 cancellata Réding, 248 20-664 cassinoides Tate, 272 20-695 Cithara Jousseaume, 237 20-653 clarki Vokes, 226 20-626 clathrata Tate, 271 20-697 conoidalis Lamarck, 247 20-663 costata Linné, 253 20-669 crassa Krauss, 242 20-658 crassa Moérch, 242 20-658 crenata Gray, 259 20-675 crenata Swainson, 258 20-674 crescentensis Weaver & Palmer, 216 20-610 davidis Link, 249 davidis Riding, 248 dechordata White, 216 delicata Perry, 250 Deniharpa Iredale, 263 doris R6ding, 255 eggs, of Harpa, 209 elegans Deshayes, 227 Eocithara Fischer, 223 Eocithara species, 23] exquisita Iredale, 263 gracilis Broderip & Sowerby, 243 grandiformis Perry, 247 gruneri ~Maltzan” Sutor, 254 Harpa Lamarck, 237 harpa Linné, 237 Harpa Pallas, 237 Harpa Réding, 237 Harpa species, 260 harpa Wood, 247 Harpalis Link, 237 Harparia Rafinesque, 237 Harpidae Bronn, 223 hilarionis Gregorio, 226 imperialis Chemnitz, 254 imperialis Lamarck, 254 jacksonensis Harris, 227 josephiniae Sacco, 257 5 key to Harpidae genera, 218 — laetifica Melvill, 254 lamellifera Tate, 235 ligata Menke, 247 loisae n. sp., Rehder, 264 Lyra Griffith & Pidgeon, 237 [20-599] [looseleaf] 20-665 20-664 20-610 20-666 20-689 20-67 1 20-603 20-635 20-623 20-635 20-689 20-659 20-663 20-670 20-653 20-653 90-653 20-653 20-676 20-663 20-653 20-653 20-623 20-626 20-670 20-670 20-627 20-673 20-616 20-670 20-647 20-663 20-690 20-653 274 Harpidae madachi Noszky, 216 major Link, 247 major Réding, 245 Marwickara Laws, 233 minor Lamarck, 242 morgani Cossmann & Pissaro, 223 multicostata Sowerby, 254 muticaeformis Martin, 230 mutica Lamarck, 224 myrmia Olsson, 259 nablium “Mart.” Morch, 249 nablium “Mart.” Sowerby, 247 narica Vredenburg, 229 neozelanica Suter, 216 nobilis Lamarck, 250 nobilis Link, 238 nobilis Réding, 238 oblonga Schumacher, 242 pachychelia Tate, 27] Palamharpa Iredale, 263 paleontological history, 212 pandura “Solander” Lightfoot, 256 pulligera Tate, 261 punctata Verco, 265 radula, 208 raricostata Risso, 227 Refluharpa Iredale, 235 H. A. Rehder 20-610 20-663 20-661 20-641 20-658 20-623 20-670 20-630 20-624 20-675 20-665 20-663 20-629 20-610 20-666 20-654 20-654 20-655 20-697 20-689 20-606 20-672 20-685 20-691 20-602 20-627 20-647 rivoliana Lesson, 259 rosea Kiener, 259 rosea Lamarck, 256 scriba Valenciennes, 259 sinuosa Stephenson, 216 solida “A. Adams” Sowerby, 242 solidula A. Adams, 242 soriensis Eames, 216 spirata Tate, 269 striatula A. Adams, 247 striata Lamarck, 249 submutica Orbigny, 228 sulcosa Tate, 269 tatei Finlay, 269 taxa, list of (Harpidae), 214 tenuis Tate, 270 testudo Donovan, 252 testudo Lightfoot, 252 tosa Aoki, 239 Trameharpa Iredale, 263 trimmeri Fleming, 216 ventricosa Lamarck, 247 ventricosa Lamarck, 25] virginalis Sowerby, 242 vulgaris Schumacher, 247 waihaoensis Laws, 233 wilsoni n. sp., Rehder, 267 Published by THE DEPARTMENT OF MOLLUSKS Delaware Museum of Natural History Greenville, Delaware 19807, U.S.A. [20-600] Index 20-675 20-675 20-672 20-675 20-610 20-658 20-658 20-610 20-695 20-663 20-665 20-628 20-695 20-695 20-608 20-696 20-668 20-668 20-655 20-689 20-610